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PROCEEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF THE
ZOOLOGICAL SOCIETY
OF LONDON.
/ f
C4
( p “ie ee
1914, pp. 491-1077,
witH 33 Puates and 129 TExtr-FIGURES.
PRINTED FOR THE SOCIETY,
SOLD AT ITS HOUSE IN REGENT’S PARK.
LONDON:
MESSRS. LONGMANS, GRHEN, AND Co,,
PATERNOSTER ROW.
DAES
lilt coeel
OF THE
COUNCIL
AND OF FLOE RS
OF THE
ZOOLOGICAL
SOCIETY
OF LONDON.
1914.
Patron.
His Masgesty THe Kina.
COUNCIL.
His Grace Tae Duke or Beprorp, K.G., F.R.S8., President.
Ricard H. Burnt, Esq.,M.A.,
Vice-President.
ALFRED H. Cocks, Esq., M.A.
THe Rr. Hon. tur Ear. or
Cromer. VPC.) G:CsB,,
CeCvivGe eK Cis. I:
Vice-President.
F. G. Dawtrey Drewirr, Esq.,
M.A., M.D.
CHarLes DrumMonp,
Treasurer.
THe Earu or Dunmore, V.C.,
M.V.O.
Sir Waurer Roper LAwreNcE
>)
Esq.,
?
Bt., G.C.L.K., Vice-President. |
Ernest W. MacBrips, Esq.,
Mes. 2Se, URS. WVice-
President.
elas. ?
|W. R. Octtvie-Grantv, Esq.
EK. G. B. Mrapr- W Avo, Esa.
Prof. Epwarp A. Muncuin,
M.A., F.R.S., Vice-President.
P. Cuatmers MircHeEyu, Esq.,
MA. DSc, LEDS Shakes
Secretary.
ABert Pam, Esq.
ApRIAN D. W. Potxock, Esa.
THe EaAru or PortsmoutH.
THe Marquess oF Suico,
F.S.A.
OLDFIELD Tomas, Esq., F.R.S.
ANTHONY H. WHuINGFIEDD,
Ksq.
Henry Woopwarp, Ksq., LL.D.,
ELRAS., Vice-President.
PRINCIPAL OFFICERS,
P. Cuatmers Mircnrnz, M.A., D.Sc., bab,
Seeretary.
Frank E. Bepparp, M.A., D.Sc.. F.R.S., Prosector.
R. I. Pocock, F.R.S., F.L.S.,
Curator of Mammals and
Tesident Superintendent of the Gardens.
D. Sera-Smivu, Curator of Birds and Inspector of Works.
Epwarp G. Bourencer, Curator of Reptiles.
Prof. H. Maxwett Lerroy, Curator of Insects.
- Henry G. Purnen, F.R.S., M.R.GS., Pathologist.
Henry G. J. Peavor, Librarian and Clerk of Publications.
Joun Barrow, Accountant.
W. H. COLE, Chief Clerk.
‘
/
LIS? OF CONTENTS.
1914, pp. 491-1077.
EXHIBITIONS AND NOTICES.
The Szcrerary. Report on the Additions to the Society’s
Menagerie during the month Olean OAS meet 2
Mr. J. Taornton Carter, F.Z.8. Exhibition of Micro-
photographs of Mitosis in the Cells of the Enamel
Organ in Dasyurus viverrinus and Trichosurus vul-
PACONGTRR ESOL Ge 2 BOR ne aOR BACHE ery URE HER OER i rere rere
Mr. R. I. Pocock, F.R.S., F.LS., F.Z.S., Curator of
Mammals. Exhibition of a white specimen of
Poumuermes: Gazelle jase. tease a. ween adceee . seers
The Rev. T. R. R. Sreppine, M.A., F.R.S., F.Z.8. Huval-
lentinia, nom. n. for Vallentinia Stebbing...............
The Secretary. Report on the Additions to the Society’s
Menagerie during the months of June, July, August,
MINOW pS\@) OUST OSTEND ee airline Bees Or cla: Aare Mer nReeeen ae
Mr. R. H. Burne, M.A., V.P.Z.8. Exhibition of some adap-
tations for the nourishment of embryos of Hlasmo-
| TEATS 0S SRE b atone Breet Ca NEE Rea ne Oe ns Ben
Mr. R. E. Savace. Exhibition of two abnormal Herrings.
Messrs. EK. Heron-Apuen, F.LS., F.Z.8., and Arrnur
Eartanp, F.R.M.S. Notice’ of Memoir on the
Foraminifera of the Kerimba Archipelago, obtained
by Dr, J. J. Simpson in the years 1907-8 ............
Page
943
943
944
944
iv
Mr. W. L. Disraxt. Notice of Repost on Rhynchota
collected by the Wollaston Expedition in Dutch New
(CUFTINry aio sat ade bobaieeates onpeaneasuse IsBtenennd snoddoonronots
‘The Sucrerary. Report on the Additions to the Society's
Menagerie during the month of October 1914 .........
The Secretary. Exhibition of a photograph of Oysters
erowing upon mangroves at Lobito Bay, Portuguese
Wiest Aitisi@a Wek ee no tcee eee soaastc na la! sam eench)aethe eee
Mr. RB. H. Burne, M.A., V.P.Z.S. Exhibition of Simu-
ibveren, ose Wikollllinisreria SINAIS, (oc cancsoccosnossconotndacaoacnadec
Mr. H. BR. Hoee, M.A., F.Z.S. Notice of Report on the
Spiders collected by the British Ornithologists’ Union
and Wollaston Expeditions in Dutch New Guinea...
Mr. Lewis H. James, B.A., F.Z.S. Notes upon the birth of
a Porpoise at the Brighton Aquarium ..................
Mr. BR. I. Pocock, F.R.S., F.L8., F.Z.8., Curator of
Mammals. Exhibition showing some new points of
difference between Pine and Beech Martens. (Text-
ined nnetss EME Oh eee haope eocecneccsostucbaroddbsaneatbceoscousdise
Mr. E. T. Newton, F.R.S., F.Z.S. Exhibition of bones of
animals showing indications of natural repair, and
some teeth of a female Physeter
Messrs. E. Heron-Auten, F.LS., F.Z.S., and Arraur
Earuann, F.R.M.S. Exhibition and Discussion on
‘“ Purpose” and ‘“ Intelligence” in the Foraminifera.
Mr. D. Sers-Suirs, F.Z.S8., Curator of Birds. Exhibition
of an Egg of the New Guinea Rifle-bird (Puilorhis
intercedens) laid in the Society's Gardens ...............
Dr. Rosrerr Broom, C.M.Z.S. Exhibition of Mammalian
Skulls showing Dental Variations ...................
Dr. Rogert Broom, C.M.Z.S. ‘Exhibition of the Skull of
a new Thecodont Reptile from South Africa. (Text-
figures 1 & 2.)
Cee decade rere emt anscececr ace ensececesee
Page
1060
1060
1061
1061
1061
1061
1062
1069
1069
33.
34.
35.
36.
38.
39.
40.
41.
PAPERS.
The Courtship-habits of the Great Crested Grebe
(Podiceps cristatus); with an addition to the Theory
of Sexual Selection. By Jurian 8. Huxtey, B.A.,
Professor of Biology in the Rice Institute, Houston,
Texcarsamm (Ilaibes ae vcz dis) SS ES cole alte aa sabe cess
On the Species of Alastor (Paralastor) Sauss., and some
other Hymenoptera of the Family Eumenide. By
R. C. L. Perxins, M.A., D.Sc., F.Z.S. (Plate I.) ...
A Report on the Fauna of the Monte Bello Islands.
By P. D. Monracur, B.A., Gonville and Caius
College, Cambridge. (Plates I—-IV.)..............0c00008
Stalk-eyed Crustaceans collected at the Monte Bello
Islands. By Mary J. Rarapun, United States
National Museum, Washington, D.C., U.S.A. (Plates
IES ip ELS) WR eae, eats} see AC a A ee
- Report on Mollusca collected at the Monte Bello
Islands. By Tom Irepate. (Text-figure 1.) .........
Cephalopoda from the Monte Bello Islands. By G. C.
RGESON, (Bo AG (Mext-teumecl.) 22 5. aeniss gar tas 2 anie='
Description of a new Lizard from the Canary Islands.
Biya eeu UME SWr ss ania cocettesaesenon: cae stcte casas
The Mechanism of Suction in the Potato Capsid Bug,
Lygus pabulinus Linn. By P. R. Awart, B.A. (Can-
tab.), D.I.C. (Lond.), Sir John Wolfe-Barry Research
Scholar, Imperial College of Science, London. (Text-
mieiees SC HOL)) warned sl See ag sors Ae Parasia Seige se scyoe
Procolophon trigoniceps, a Cotylosaurian Reptile from
South Africa. By D. M.S. Warson, M.S8c., F.Z.8.,
Lecturer on Vertebrate Paleontology in University
College, London. (Plates I-III., and Text-figures
1] thio) ee On cea ee eA mabe Nel geiko Ors Ea re See
P
Lon
age
491
4)
63
(or)
bo
Or
6
6
6
6
ay)
65
17
81
45.
44,
46.
48,
AQ,
Vi
. he Deinocephalia, an Order of Mammal-like Reptiles.
By D. M.S. Watson, M.Sc., F.Z.S., Lecturer on Verte-
brate Paleontology in University College, London.
(Plates IV., V., and Text-figures AKSS Meas Sones sane
Diagnoses of New Genera and Species of Zonitide from
Equatorial Africa. By H. B. Prusrox, F.ZS.
(U2) biitsiof EIB) ieee ssaanpe daddeuasorabnrpchoéaabendoncbooco pon
On a second Collection of Batrachians and Reptiles
made by Dr. H. G. F. Spurrell, F.Z.5., in the Choco,
Colombia. By G. A. BounencEr, F.R.S., F.Z.8.
(LEE Ri ese ea Oe Aso neMeRanSanRBe onausebopaconaobecseagnds00<
. Zoological Results of the Third Tanganyika Expedition,
conducted by Dr. W. A. Cunnington, 1904-1905,.—-
Report on the Parasitic Hucopepoda. By WinuiAM
A. Cunnineton, M.A., Ph.D., F.Z.S. (Plate L., and
Mexbomoume ge). so oie chats ea noe ae once eee eee
The Marine Fauna of British Kast Africa and Zanzibar,
from Collections made by Cyril Crossland, M.A.,
B.Sc., F.Z.8., in the Years 1901-1902. Bryozoa
Cyclostomata, Ctenostomata, and Endoprocta. By
Artur Wm. Waters, F.L.S., F.G.8. (Plates I-IV.,
and Text-figure 1.)
Cer ew meee ete s eect eens ese snc es occ ease ress e
. Contributions to the Anatomy and Systematic Arrange-
ment of the Cestoidea.—XIV. On a new Species of
thabdometra, and on the Paruterine Organ in Oti-
ditema. By Frank E. Bepparp, M.A., D.Sc., F.R.S.,
F.Z.8., Prosector to the Society. (Text-figures 1-11.)
On the Facial Vibrisse of Mammalia. By R. I.
Pocock, F.R.S., F.L.S., F.Z.8., Curator of Mammals.
(Text-figures 1-13.)
\c)e/sieleie:p/ei¢isis/eia\e}e je cjaiylejeieiejajsle/elalainie)syele?aicievetate
On the Feet and other External Features of the Canidse
and Urside. By R. I. Pococx, F.R.S., F.L.S., F.ZS.,
Curator of Mammals. (Text-figures 1-13.)
- A remarkable new Cirripede from the Chalk of Surrey
and Hertfordshire. By Tuomas H, Wiruers, F.G.S.
(Plate L., and 'Text-fignre tS)
#018 6) 0,)6)9\0\(0. 0/40 an e)els\e sisneldvale itinerary,
Page
787
813
819
831
859
889
51.
52.
53.
o4.
55.
Vil
Polycheta from the N.H. Pacific: The Chetopteride.
With an Account of the Phenomenon of Asexual
Reproduction in Phyllochetopterus and the Description
of Two new Species of Cheetopteride from the Atlantic.
By F. A. Porrs, M.A., Fellow of Trinity Hall,
Cambridge, and Balfour Student of the University.
(Plates I1.—VI., and Text-figures 1-i3.)..................
Broomia perplexa, gen. et sp. n., a Fossil Reptile from
South Africa. By D. M.S. Watson, M.S8c., F.Z.8.,
Lecturer on Vertebrate Paleontology in University
College, London. (Plates VI., and Text-figures 1-5.)
Hunotosaurus africanus Seeley, and the Ancestry of the
Chelonia. By D, M..S. Watson, MSc, E.Z:s.,
Lecturer on Vertebrate Paleontology in University
College, London. (Plate VII., and Text-figure 1.)...
Notes on some Carnivorous Therapsids. By D. M.5.
Watson, M.Sc., F.Z.8., Lecturer on Vertebrate Pale
ontology in University College, London. (Text-
SCOTT) 11a) ae a ee NSO ree tree RA a Nc
Contributions to the Anatomy and Systematic Arrange-
ment of the Cestoidea—XV. On a new Genus and
Species of the Family Acoleide. By Frank E.
Brpparp, M.A., D.Sc., F.R.S., F.Z.S., Prosector to
mmepsocte iyi, (hext- Mo umesil Oe oe css s sac sje oon ses
Page
955
995
1011
Dae
ie Oe F ‘ ‘en
j i _ ne Bs
: sh hae in brisie: a,
oh
iA BECCA. Gist
OF THE
CONTRIBUTORS,
With References to the several Articles contributed by each.
(1914, pp. 491-1077.)
Page
Auten, HE. Heron-. See Heron- Aen, E.
Awatt, P. R., B.A. (Cantab.), D.I.C. (Lond.).
The Mechanism of Suction in the Potato Capsid Bag,
Lygus pabulinus Linn. (Text-figures 1-29.)............... 685
Bepparp, Frank E., M.A., D.Sc., F.R.S., F.Z.8., Prosector
to the Society.
Contributions to the Anatomy and Systematic Arrange-
ment of the Cestoidea :—
XIV. On a new Species of Rhabdometra, and on the
Paruterine Organ in Otaditenia. (Text-figures 1-11.) ... 859
XV. On a new Genus and Species of the Family
meorerdeewe (Nexb-tommes 192) Oeil ecccsecseesseckces: 1039
BovuLencer, Georce A., LL.D., F.R.S., F.Z.S.
On a second Collection of Batrachians and Reptiles
made by Dr. H. G. F. Spurrell, F.Z.S., in the Choco,
Colermiiraien (eleiiesile MP eee ee s.r hs eles cac endo ee 813
x
Page
Broom, Rosert, M.D., C.M.Z.S8.
Exhibition of Mammalian Skulls showing Dental
ICE Le RIUTOVa iBone “unre Nar nnena ne acbint see bene ane adie eM ares ous 1071
Exhibition of the Skull of a new type of Thecodont
Reptile (Youngina capensis) from the Upper Permian
Beds of South Africa. (Text-figures 1 & 2.) ............... 1072
Burne, Ricwarp H., M.A., V.P.Z.S.
Exhibition of some adaptations for the nourishment
of the embryos of Elasmobranchs ..............0...0:000000: 1059
Exhibition of Simulacra of Molluscan and other
PED S mere Meet sedi scab adae clive anaes eho te Ae eae 1061
Carter, J. THorNToN, F.Z.S.
Exhibition of Microphotographs of Mitosis in the
Cells of the Enamel Organ in Dasyurus viverrinus and
Driehoeumus wulpoowla 234i ae Oe, ee 943
Cunnineton, Witiiam A., M.A., Ph.D., F.Z.S.
Zoological Results of the Third Tanganyika Expedi-
tion, conducted by Dr. W. A. Cunnington, 1904-1905,.—
Report on the Parasitic Eucopepoda. (Plate I., and
‘bes Feino (Uae 1) AN ee em aren De ML oo el 819
Distant, WiLuiAM L.
Notice of Report on the Rhynchota collected by the
Wollaston Expedition in Dutch New Guinea............... 1060
Earuannd, ArtuuR, See Heron-Auven, E.
x1
Heron-Atien, Epwarp, F.L.S., F.Z.8., and Haranp,
Artruur, F.R.M.S.
Notice of Memoir on the Foraminifera of the Kerimba
Archipelago obtained by Dr. J. J. Simpson in the
years 1907-8
See eee CC CC tC a
Exhibition and Discussion on ‘‘ Purpose” and “ In-
telligence”” shown by the Arenaceous Foraminifera in
the construction of their tests
HocewriNnY i, MAL HZS.
Notice of Report on the Spiders collected by the
British Ornithologists’ Union and Wollaston Expeditions
in Dutch New Guinea
Houxtey, Juxian §., B.A.
The Courtship-habits of the Great Crested Grebe
(Podiceps cristatus) ; with an addition to the Theory of
SexualySelections, (Plates Wd UN) n 00. \.ccbcseme sence =
IREDALE, Tom.
Report on Mollusca collected at the Monte Bello
Islands. (Text-figure 1.) ..........:.cceeeeteee eects enene tens
JAMES, Lewis H., B.A., F.Z.8.
Notes upon the birth of a Porpoise at the Brighton
JA ORURN GTO | 152 Alo kenan du asbag coe) pcoche AA HBHROeO nun oh age Searing
Leurs, Dr. Pu.
Description of a new Lizard from the Canary
TISTIIGIS | GlectkeoenSehon doc npeorsdete och au. cc no co pALHCp ADSM oan pno
Page
1060
1069
1061
491
665
1061
X11
MircuHecy, P. Caatmers, M.A., D.Sc., LL.D., F.R.S., F.Z.8.,
Secretary to the Society.
Report on the Additions to the Society’s Menagerie
dumne themonbh of Mays OA yee) ee ceehn.care seer nere
Report on the Additions to the Society’s Menagerie
during the months of June, July, August, and September,
SU et nee se Je cto etal Oke eh UR mI is 2 ot cig aiallt miele
Report on the Additions to the Society's Menagerie
gurine thewmonth ot October l Qa eee he ceacecse scree
Exhibition of a photograph showing Oysters growing
upon mangroves at Lobito Bay, Portuguese West
JANA GUAGE Wp oasee aa eters. 16 Se Tee eh hs te SA a a
MownracuE, P. D., B.A.
A Report on the Fauna of the Monte Bello Islands.
(ERG eS BU IEV i. iy ieccne meeri. ation (tre ise Ue cue par ea
Newron, Epwin T., F.R.S.,. F.Z.S.
Exhibition of bones of animals showing indications of
natural repair, and some teeth of a female Physeter ......
Perkins, Ropert C. L., M.A., D.Sc., F.Z.8.
On the Species of Alastor (Paralastor) Sauss., and
some other Hymenoptera of the Family Eumenide.
(ESE ie Rk) RIE AP a eee eee oct g Da IRD Ze URN ity Ine Wc
Pocock, Reerarp I; F.R.S., F.LS., F.Z.8., Curator of
Mammals.
On the Facial Vibrissee of Mammalia. (Text-figures
1-13.)
On the Feet and other External Features of the
Canidee and Urside. (Text-figures L=13:)
Exhibition, on behalf of Major C. P. Bradshaw, of a
white specimen of Sémmering’s Gazelle
Exhibition showing some new points of difference
between the Pine Marten (Martes martes) and the Beech
Marten (Martes foina). (Text-figures 1-4.)
Page
1069
563
889
913
944
xi
Ronis Ac. VE AG
Polycheta from the N.E. Pacific: the Chetopteride.
With an Account of the Phenomenon of Asexual
Reproduction in Phyllochetopterus and the Description
of Two new Species of Cheetopteride from the Atlantic.
(Blabes We Vile ame Pext- tomes 30) a sess aa. vee:
Preston, Hucu B., F.Z:S.
Diagnoses of New Genera and Species of Zonitidee
omemauatoartale Amica. (Plates ill) oo se scecc0..5-
Rarupun, Miss Mary J.
Stalk-eyed Crustaceans collected at the Monte Bello
sland seuaG@ilaites: Uc ds MAC) Sie. .erteasvieeeencl se eladed. cece
Rosson, G. C., B.A.
Cephalopoda from the Monte Bello Islands. (Text-
i@inine dl py Seeea dee nhe een riant Ome se eRe Ebene Seat onto anaes
Savace, R. E.
Exhibition of two abnormal Herrings, taken by trawl
in the North Sea
Seem em eee men enter ewe wees eee e reese eee e ee ee eee sere
Sura-Smrra, Davin, F.Z.8., Curator of Birds.
Exhibition of an Egg of the New Guinea Rifle-bird
(Piilorhis intercedens) laid in the Society’s Gardens ......
Srepernc, The Rev. THomas R. R., M.A., F.B.S., F.LS.,
E.ZS.
Serene ree
Huweallentinia, nom, n. for Vallentinia Stebbing
955
787
653
677
1060
X1V
WATERS, ARTHUR W., PLS. BGS.
The Marine Fauna of British East Africa and
Zanzibar, from Collections made by Cyril Crossland,
M.A., B.Sc., F.Z.S., in the Years 1901-1902. Bryozoa—
Cyclostomata, Ctenostomata, and Endoprocta. (Plates
SII chats Neprareimeiiaey NS) Best ecooneccdeasspocedneeapoacacca:
Watson, Davin M. §., M.Sc., F.Z.S.
Procolophon trigoniceps, a Cotylosaurian Reptile from
South Africa. (Plates I—ITI., and Text-figures 1-5.) ...
The Deinocephalia, an Order of Mammal-like Reptiles.
(eibneay HW, Wop eumal Wescoctnlegnaaess WIS) oascaonoceoseavsedion
Broomia perplera, gen. et sp. n., a Fossil Reptile from
South Africa. (Plate VI., and Text-figures 1-5.)
Eunotosaurus africanus Seeley, and the Ancestry of
the Chelonia. (Plate VII., and Text-figure 1.)
Notes on some Carnivorous Therapsids. (Text-
figures 1-7.)
Wrruers, Tuomas H., F.G.S.
A remarkable new Cirripede from the Chalk of Surrey
and Hertfordshire. (Plate I, and Text-figure 1.)
831
INDEX.
1914.—Pages 491-1077.
[New names in clarendon type.
Systematic references in italics.
(z.8.1.) indicates additions to the Society’s Menagerie. |
Abispa meade-waldoensis, sp. n., |
623.
Ablepharus muelleri: ethology, 641.
Acanthopleura, 669.
gemmata, 666, 668.
spinosa, 666, 668.
Acmea saccharina, var., 666, 670.
Acridium maculicollis, 648.
Actea affinis, 688.
—— glandifera, sp. n. (PI. I. fig. 5),
658.
Acte@odes affinis, 658.
Actumnus setifer, 660.
Adelosaurus, gen. n., 1009.
huxleyi: structure (Fig. 5), 1008.
JEpyprymnus rufescens: facial
brissz (Fig. 3), 893.
Africarion concavospira, sp. n.
(Pl. II. fig. 20), 787.
—— copiosa, sp. n. (Pl. Il. fig. 18),
788.
—— kagambahensis, sp. n. (PI. II.
fig. 18), 789.
—— kiduhaensis, sp. n. (Pl. I.
fig. 16), 788.
marsabitensis, sp. n. (Pl. IT.
fig. 14), 788.
orestias, sp. n. (Pl. II. fig. 15),
788.
Vi-
—— oscitans, sp. n. (Pl, II. fig. 4),
788.
—— spatiosa, sp. n. (Pl. II. fig. 17),
789.
Africarion tenebrosa, sp. n. (PI. IL
fig. 19), 789.
Alastor, 563.
Alectrion suturalis, 667.
Alopex lagopus: feet: rhinarium:
facial vibrissee (Fig. 7), 923.
Alpheus bucephalus, 654:
—— edwardsti, 6)4.
Amalda elongata, 667.
Amathia dichotoma, 848.
distans, 848. '
-—— lendigera (Pl. IV. figs. 3, 4), 847,
848.
semiconvoluta, 847.
vidovici (Pl. IV. figs. 1, 2), 848.
Ammotragus lervia: facial vibrisse, —
911.
Amsacta marginata, 645.
Amyna. octo, 647.
spilonota, 647.
Anastomus oscitans (Z. s. L.), 943,
Anatomy. See Srrucrure.
Anchistus inermis, 656.
Ancilla elongata, 667.
Anoa depressicornis (z. 8. u.), 1057.
Anolis palmeri, 814.
Anthela pudica, 646.
Anthus australis montebelli, 635.
Antigona tiara, 666, 668.
Antilope cervicapra: facial vibrisse,
911.
Anumeta zuboides, sp. nu. (Pl. I.
fis, 12), O47,
xV1
Aporus sp., 649.
cingulatus, 649.
ARACHNIDA :
Aranee from Dutch New Guinea
(see ‘ Transactions ’).
Aramus giganteus (z. 8s. L.), 1058.
Arca fusca, 666.
Archimantis brunneriana, 648.
Arctocephalus pusillus: facial vibrissee,
901.
Arctognathus
1028.
Arctops willistoni, gen. et sp. u.:
structure (Figs. 3, 4), 1027.
Arcularia suturalis, 657.
Artamus leucorhynchus harterti, 654.
Asanadopsis mgobergi, 65).
curvimola: structure,
Ascopodaria gracilis, 85.
Atelopus spurrelli, sp. n. (Pl. I.
fig. 1), 815.
Atergatis floridus, 657.
ocyroé, 657.
Atherura africana:
(Pig. 9), 903.
Atractaspis irregularis (z. s.u.), 1059.
Aves:
from Monte Bello Islands: syste-
matic: ethology, 632; Podiceps
cristatus: courtship habits, 491 ;
Ptilorhis
1070.
Axis axis: facial vibrisse (Fig. 12),
909.
facial yvibrissze
intercedens: ethology,
Barentsia gracilis, 855,
Barracnuta :
from Choco, Colombia: systematic,
813.
Bauria cynops:
(Fig. 1), 1021,
Beania intermedia, 856.
Belenois java, 644.
—— teutonica, 644.
cayennensis
structure
(skull)
Belonopterus
1050.
Bembex (?) variabilis, 649.
Blayneyella, gen. n., 799.
—— kisengiensis,
fiz, 21), 799.
(z. S. L.),
(Qt I
sp. n.
INDEX.
Blayneyella microspiralis, sp. n.
(Pl. I. fig. 18), 800.
percivali, sp. n. (PI. I. fiz. 19),
799.
purpureocincta, sp.n. (Pl. I.
fig. 20), 79).
Boa imperator, 815.
Bos frontalis: facial vibrissee, 911.
Boselaphus tragocamelus: facial vi-
brissee, 911.
Bostrychopsis jesuitus, 647.
Bowerhankia pustulosa, 851.
Bradypus tridactylus: facial vibrissze,
893.
Broomia perplexa, gen. et sp. n.:
structure (Pl. V. figs. 1-4), 995.
Lruchigavia longirostris, 637.
nove-hollandie longirostris, 637.
Bryozoa. See Ponyzoa.
Bufo coniferus, 813.
Bu'laria columellaris, 667.
Burungaélla, gen. n., 797.
buhambaénsis, sp. n. (Pl. J.
fig. 16), 798.
imperforata, sp. n. (Pl. I.
fig. 15), 798.
mutandana, sp. n. (Pl. I.
fic. 14), 798.
oscitans, sp. n. (Pl. I. fig. 17),
Tote
Buskia nitens, 854.
Callithrix jacchus: facial vibrisse, 898.
Camelus bactrianus: facial Vibrissxe, |
SU
dromedarius: facial vibrissee, $11.
Cancer dama, 662.
Canis anthus: feet (Fig. 4), 918.
mesomelas: feet: rhinarium :
facial vibrissee (Fig. 7), 901 ; (Fig. 4)
920.
pallipes: feet (Fig. 1), 915.
— sclateri: feet: rhinarium : facial
vibrissx (Fig. 5), 920,
Cardita incrassata, 666.
Cardium dupuchense, 666.
——— unedo, 666, 667.
Curpilodes ruber, 657.
Castor canadensis (z. 8. 1.), 1058,
INDEX.
Cavia rufescens: facial vibrisse
(Fig 10), 903.
Centetes ecaudatus :
(Fig. 4), 898.
Cephalophus coronatus: facial vibriss
(Fig. 13), 911.
Cephalopyrus flammiceps
1057.
Cercopithecus: facial vibrisse, 898.
Cerdocyon microtis: feet: rhinarium :
facial vibrissee (Fig. 5), 920.
Cerithium fasciatum, 606.
Certhiola flaveola (z.s.u.), 1058.
Cervus elaphus: facial vibrisse, 910.
eldi: facial vibrisse, 909; (z.s.L.),
1057.
Chetopteride: structure: development
(Pls. I-VI. figs. 1-13; Figs. 1-13),
955.
Chetopterus variopedatus: structure:
development (Figs. 1, 2), 961, 978.
Charadrius geoffroyt, 637.
— mongolus, 637. :
ruficapillus tormenti, 637.
Chelonia: structure, 1011.
Chelydra rossignont, 814.
Chinchilla lanigera: facial vibrisse,
903.
Chiton cunninghamt, 669.
Chiamys lentiginosus, var., 666.
radula, 666.
—— squamosus, var., 666.
Chloridea arnugera, 646.
Cheroichthys valenciennet, 650,
Cheropotamus : facial vibrisse, 908.
Chrysochloris asiatica: dental varia-
tion, 1071.
— hottentota :
LOGE
Chrysococcyx basalis wyndhami, 633.
Cicinnurus regius (z. 8. u.), 1059.
Cinnyris chalybeus (z. s. u.), 1058.
Cirphis abdominalis, 647.
Cissa chinensis (z. s. u.), 1060.
Clupea harengus: abnormal, 1060.
Coccinella transversalis, 647.
Celogenys paca: facial vibrisse, 903.
Coendu bicolor (z. s. u.), 1057.
prehensilis: facial vibrisse, 903.
facial vibrissa
(Z. 8. U.),
dental variation,
iF
XVll
Collusa sp. ?, 646.
Coluber arizone (z. s. u.), 1057.
——- corais, 815.
Connochxtes albojubatus (z, s. L.),
1059,
Conurus canicularis (z. s. L.), 948.
Conus anemone, 667, 675.
maculosus, 6795.
Cormocephalus turneri, 650.
Coryphistes cyanopterus, 648.
Coua ruficeps (2. 8. u.), 1057.
Crangon bucephalus, var., 654.
edwardsii, 654.
Cricetomys gambianus: facial vibrissee,
902.
Crisia circinata, sp. n. (Pl. I.
fics. 7-9), 840.
—— denticulata (Pl. IV. fig. 5),
837.
elongata (Pls. I. figs. 3, 4; IV.
fig. 6), 838.
inflata, sp. n. (Pl. I. figs. 1, 2),
839.
recurva, 839.
sertularoides (Pl. I. figs. 5, 6),
839.
Crotaphytus collaris (z.s. u.), 1058.
CRUSTACEA :
from Monte Bello Islands: syste-
matic: ethology, 653.
Parasitic Kucopepoda: Tanganyika :
River Nile: systematic, 819.
Cirripedia: systematic: structure,
945,
Ctenostomata, 831.
Cuon primevus: feet: rhinarium :
facial vibrissx (Fig. 2), 916.
Cyclostomata, 831.
Cylindrecium giganteum, 854.
Cymatium aquatile, 666.
Cyprea caputserpentis, 667.
caurica, 667.
—— cylindrica, 667.
——- erosa, 667.
errones, 667.
moneta, 567.
Cyrtacanthus guttulosa, 648.
Cystophora cristata: facial
Gis. 7); 901; (@ s.u.), 943.
vibrisse
XVIl1
CytTroLoGy:
Manmarta: Dasyurus: Trichosurus: |
enamel-cells, 9-43.
Dama dama: facial vibrisse, 910.
Damaliscus albifrons (z. s. u.), 1057.
Danais chrysippus f. petilia, 644.
Dardanus megistos, 656.
Dasyprocta columbiana: facial vibrisse,
903.
Dasyurus viverrinus: facial vibrisse,
891; cytology, 943.
Daubentonia (Chiromys) madagascari-
ensis: facial vibrissee (Fig. 6), 898.
Deinocephalia: structure (Pls. 1V., V.;
Figs. 1-18), 749.
Demiegretta sacra: ethology, 638.
Dendrohyrax dorsalis: facial vibrisse,
906.
Dendrolagus ursinus; facial vibrissee |
o
(Fig. 3), 895.
Dermestes cadavorinus, 647.
Deuterosaurus, 774.
DEVELOPMENT:
Mammatia: Cetacea: Phoczna cow-
munis, 1061.
Verminra: Phyllochetopterus, 972.
Diademodon: structure (Figs. 3, 4), |
1028.
Didelpbys azar:
(Fig. 1), 891.
Dimetrodon: structure (Fig. 4), 1026.
Diphyllodes hunsteini (z. s. u.), 1060.
Dolichotis salinicola :
903.
Drymobius boddaertii, 815.
facial
Echidna hystrix: facial vibrisse, 891.
Ectopatria aspera, 646.
Egernia striolata (z. s. u.), 1058.
Elaps spurrelli, sp. n.
fig. 3), 817.
Elgonella, gen. n., 795.
—— angustior, sp. n. (Pl. IT. fig. 6), |
797.
brunnea, sp n. (PI. II. fig. 11), |
796.
—— discolorata, sp.n, (Pl. IL. fig.7),
796.
vibrissxe |
facial yvibrisse, |
(CEL elale |
INDEX.
Elgenella euloteformis, sp. n.
(BIS ties9);) 796:
——- flavidula, sp. n. (Pl. Il. fig")
796.
oribates, sp. n. (PI. II. fig. 8),
197.
robini, sp. n. (P1. Il. fig. 10),
797.
sobrina, sp. n. (PI. Il. fig. 12);
797.
Allipsidron inguinata, 648.
Endoprocta, 831.
Entalophora deflexa, 840.
——— wasinensis, nom. un.
figs. 1-4, 9: Fig. 1), 840.
Eosuchia, subord. nov., 1077..
Epactrothynnus productus, 649.
Ephbthianura albifrons (z. s. u.), 105°.
——— tricolor distincta, 634.
Hphutomorpha modesta, 649.
morosa, 649.
(eal Iu
| Kpicrates cenchris, 815.
Epimys norvegicus: facial vibrisse
(Fig. 8), 901.
Eptesicus pumilus : ethology, 631.
Equide: facial vibrissx, 906.
| KHremiornis carteri assimilis, 634.
Erethizon dorsatum: facial yibrissz
(Fig. 10), 903.
Hrinaceus europexus: facial vibrissee
(Big. 4), 895.
ErnouoGcy:
Aves: Podiceps cristatus: court-
ship-habits, 491; Ptilorhis inter-
cedens, 1070.
Insecta: Paralastor, 563.
Mouuusca: Ostrea, 1061,
VermipEa: Mesocheetopterus, 938 3
Phyllocheetopterus, 980.
Prorozoa: Foraminifera, 1069,
Eublemma dubia, 647.
Huchelus atratus, 666. j
Eulima montagueana, sp. n.
(Fig. 1), 672.
EHunotosaurus africanus : structure
(Pl. VII.; Fig. 1), 1015.
Euphractus villosus: facial yibrissz
(Fig. 4), 898.
Hupoda geoffroyi, 637.
INDEX.
Eupoda mongolus, 637.
Euproctis chionitis, 646.
Eutamias quadrivittatus :
brissx, 903.
Euvallentinia, nom. n. for Vadlen-
dinia, 944.
Euxoa radians, C46.
facial vi-
Falloonella, gen. n., 809.
—— exquisita, sp. n. (Pl.I. fig. 9),
809.
—— —— gudei, subsp. n. (PI. I,
fig. 10), 809.
larogiensis, sp.n. (PI. I. fig. 11),
809.
Farrella atlantica (P\. LV. fig. 9), 852.
gigantea, 854.
Filisparsa irregularis, 844.
— tubulosa, 842.
Foraminifera: ethology, 1069:
Kerimba Archipelago (see ‘ Trans-
actions’).
Fragum unedo, 667.
from
Galago crassicaudata: facial vibrissze
(Fig. 6), 898.
Gampsonyx swainsoni (z. 8. L.), 1058.
Gazella semmeringi: variation, 944.
Gehyra variegata : ethology, 640.
Gelasimus forcipatus, 661.
GEOGRAPHICAL:
Fauna of Monte Bello Islands, 625,
653, 665, 677.
ReprtuiA: Colombia: systematic,
813; South Africa (fossil), 739,
749, 995, 1021; Lacerta: Canary
Islands, 681.
Barracura : Colombia: systematic,
813.
Insecta: Hymeuoptera : Paralastor :
Australia, 563; Rhynchota: Dutch
New Guinea (see ‘ Transactions’).
Anracuyipa: Aranese: Dutch New
Guinea (see ‘ Transactions *).
Crustacna: Parasitic Eucopepoda:
Tanganyika (River Nile), 819;
Cirripedes from Chalk of Surrey |
and Herts, 945. |
ie
GuoGRAPHICGAL (con.):
Moutuvusca: Zonitide : Equatorial
Africa, 787.
Ponyzoa: B.E. Africa: Zanzibar
831.
VermipEA: Chetopteride ; N.E,
Pacific: Atlantic: British Waters
95d.
Protozoa: Foraminifera ; Kerimba
Archipelago (see ‘ Transactions’).
Geopelia humeralis headlandi, 632.
Geranoaétus imelanoleucus (z. s. L.),
1060. 3
Giraffa camelopardalis (z. s. u.), 1059.
Glis glis: facial vibrissxe (Fig. 8), 902.
Glyptoxanthus cymbifer, sp. n.
(Pls. I., IL., figs. 6,7), 658.
Gobius phalena, 650.
Gonitis subulifera, 646.
Gonocephalum meyricki, 647.
Gonodactylus chiragra,
664.
Gorgonops: structure (Fig. 6), 1031,
1034.
Graninodes oceliata, 646.
Gudeeélla aranea, sp. n. (Pl. III.
fig. 20), 789.
var. smithit,
bartaensis, sp. n. (Pl. III.
fig. 6), 790.
consobrina, sp. n. (Pl. IIL.
fig. 9), 794.
consueta, sp. n. (PI. III. fig. 18),
790.
—— densesculpta, sp. n. (Pl. III.
fig. 14), 791.
—— elgonensis,
fig. 19), 791.
—— gerstenbrandti, sp. n. (Pi. IIT.
fig. 7), 792.
—— inclinans, sp. nu. (Pl. II1. figs. 1,
8), 792.
GO, ws Ce, MOE
inflata, sp. n. (Pl, III. fig. 21),
792.
iridescens, sp. n. (Pl. III.
fig. 15), 793.
kampalaensis, sp. n. (Pl. III,
fig. 10), 790.
marsabitensis, sp. u. (PI. III.
fig. 22), 798.
xX
Gudeélla masakaénsis,
(PL IIL. fig. 16), 798.
— mime, sp, n. (Pl. III. fig. 23). 791-
mukandaensis, sp. n. (Pl. Il.
fig. 11), 794.
——, mutandana,
(Pl. III. fig. 12), 794. |
multistriata, sp. n. (Pl. III.
fig. 3), 794.
nemorum,
fig, 24), 794.
pallidior, sp, u. (Pl. III. fig. 5),
795.
tribulationis, sp. n. (Pl. III,
fic, 4), 795.
urguessensis, sp. n. (Pl. III.
fig. 17), 790.
usitata, sp. n. (Pl. III. fig. 2):
791.
woodhousei, sp. n. (Pl. III.
fiz. 13), 792.
Spey, tl.
var. n.
(Pl. IIT. |
sp. n.
Hematopus longirostris, 637.
unicolor ophthalmicus, 637.
Haleyon sanctus westralasianus, 633.
sordidus melvillensis, 633.
—— westralasianus, 633.
Haliatius leucogaster :
TIL), 638.
Haliastur girrenera: ethology, 6389.
facial vibrissz, |
ethology (Pl. |
Halicherus grypus:
901.
Haliotis squamata, 666.
varia, 666.
Halobates, 649.
Helarctos malayanus: feet: rhinarium
(Fig. 15), 936, 937.
Henicospilus sp., 649.
Hermatobatodes sp., 649.
Heteronota binoet (PI. L. figs.
ethology, 640, 641.
Hinula australis, 642,
tenuis, 642.
Hippopotamus amphibius: facial vi-
brissze, 908.
Hippuraria verticillata, 858.
Holochila heathii aerata, subsp.
1-3) :
n., 640,
INDEX.
Homalocranium nigrum, sp. n.
(Pl. II. fig. 2), 816.
Hoplodactylus pacificus (z. s. u.), 1059.
Hornera violacea, var. tubulosa, 842.
Fuenia proteus, 661.
Hyastenus oryx, 661.
Hydrocherus hydrocherus: facial vi-
brissee (Fig. 9), 904.
Hydroprogne tschegrava strenua, 636.
Hylodes ramiformis, 813.
Hyrtacus eutrachelia, 648.
Hystrix: facial vibrisse, 903.
Idmonea interjuncta (Pl. II. fig. 5),
846.
—- wregularis, 843.
milneana, 844.
-— pedleyi, 846.
pulcherrima, 846.
—— radians (Pl. II. figs. 6-8), 844.
—— —, var. erecta (Pl. II. fig. 10),
844,
Iguana tuberculata, 815.
Insecta:
Monte Bello Islands, systematic:
ethology, 643.
Hymenoptera: Paralastor: syste-
matic, ethology, 563.
Rhynchota: structure (suction -
mechanism), 685; from Dutch
New Guinea (see ‘Transactions ’).
Irena turcosa (z, 8. t.), 1059.
Isodon novitius, 647.
Lsoodon barrowensis, 631.
Jaculus jaculus?: facial vibrissee, 903.
orientalis ; facial vibrisszx, 902.
Lacerta czesaris, sp. n., 681.
Lachesis schlegelii, 817.
Lagenipora socialis, 856.
Lagorchestes conspicillatus : ethology,
630.
Lagostomus trichodactylus: facial yi-
brissze, 905.
Lagothrix lagotrichia: facial vibrisse,
898.
INDEX.
Lalage tricolor (z. 8. u.), 1058.
Lama huanacos : facial vibrissee, 911.
911;
vicugna: facial vibrissie,
(a. 8. u.), 1057.
Lambrus (Rhinolambrus)
663.
Lamiasaurus newtoni, gen. et
sp. n., structure (Figs. 8, 9), 750,
768.
Larogiella, gen. n., 800.
—— angulifera, sp. n. (PI. II. fig. 1),
800.
—- fonticula, sp. n. (Pl. II. fig. 3),
801.
kombaensis, sp. n. (PI. IT.
fig. 2), 801.
malasanjiensis, sp. n. (Pl. I.
fig. 25), 801.
venatoris, sp. n. (Pl. I. fig. £2)
800.
Ledoulxia adjacens, sp. n. (Pi. I-
fic. 8), 808.
crassiplicata, sp. n. (Pl. I.
fig. 1), 806.
—— decussata, sp. n. (Pl. L. fiz. 7),
806.
—— elgonensis, sp. n. (Pl. L. fig. 4).
807.
—— eussoéngis, sp, n. (Pl. I. fig. 2),
807.
—— jingaénsis, sp. n. (PI. L. fie. 6),
807.
—-— levistriata nyeriensis, var.
n. (Pl. I. fig. 3), 808.
—— marsabitensis, sp. n. (Pl. I.
fic. 5), 808.
Lemur eatta: facial vibrissze, 898.
fulvus: facial vibrissse. £98.
mongos: facial vibrissx, 898.
—-yarius: facial vibrisse (Fig. 6),
898.
Leontocebus leoninus ;
(Fig. 6), 898.
Lepidoblepharis intermedius,
Span (alate 2), Cit.
Lepralia poissonti, 856.
Leptophis brevior, sp. n. (PI. II.
fig. 1), 815.
pelagicus,
facial vibrissse
XX1
Lerneocera, 820, 822.
-—— diceracephala, sp. n. (Pl. I.
figs. 1-3), S24,
haplocephala, sp. n. (Pl. I.
figs. 4-7), 826.
temnocephala, sp. n. (Pl. I.
fies. 8, 9), 827.
Liasis childreni, 643.
Lima fragilis, 666.
lima, 666.
multicostata, 666.
Lophactea granulosa, 658.
_ Lophognathus gilberti, 641.
Loxosoma singulare, 855.
Lycaon pictus: feet: rhinarium : facial
vibrisse (Fig. 3), 917.
Lychas variatus, 650.
Lycosuchus: structure (Fig. 7), 1035.
Lygosoma bipes, 641.
-—— isolepis,: ethology, 642.
lesueurti: ethology, 642.
moco (Z. 8. u.), 1059.
tenue (z. Ss. u.), 1059.
Lygus pabulinus: structure (suction-
mechanism) (Figs. 1-29), 685.
Maerocorax fuscicapillus (z. s. u.), 1059.
Maeropus bennetti :
895.
billardieri: fazial vibrissx, 893.
Matleus malleus, 666.
MAMMALIA :
Facial vibrissze, 889.
Monte Bello Islands: systematic:
ethology, 630.
Canidz and Ursidz: systematic :
structure (feet: rhinaria: facial
vibrissz), 915.
Chrysochloris asiatica : C. hottentota :
dental variation, 1071.
Gazella scimmeringi: variation, 944.
Martes martes: M. foina : structure,
1062.
Phascolaretus cimereus : dental yaria-
tion, L071.
Phocsena communis:
1061,
facial vibrissze,
development,
XXi1
MawManra (com.) =
Trichosurus vulpecula : dental varia-
tion, 1071.
Manis tricuspis: facial vibrisse (Fig. 4),
895.
Marmosa elegans: facial vibrisse, 891.
Martes foina: structure (Figs. 2, 3),
1062.
— martes: structure (Figs. 1, 38),
1062.
Mazama tema?: facial vibrissee, 908.
Megapodius duperreyi (4 s. L.). 1059.
Meles meles: facial vibrissx, 901.
Melicleptria albivenata, sp. n.
(Pl. I. fig. 11), 646.
neurias, O46.
Melursus ursinus: feet:
(Figs. 12, 15), 933, 937.
Mesocheetopterus, gen. n., 957, 967.
—— minuta, sp. n.: structure (Pls.
IL., ITI. figs. 7, 8; Figs. 4, 5), 958, 963.
taylori, sp. n.: ethology: struc-
rhinarium
ture (Pls, I., III. figs. 5-6, 9; Figs. |
1, 5), 957, 958.
Metapeneus monoceros, 653.
stridulans, 693.
Microgomphodon oligocynus, structure
(skull) (Fig. 2), 1028.
Microtus sandayensis (z. s. t.), 1058.
Mictotragus arachne, 647.
Mictyris longicarpus, 661.
Mikenoélla, gen. n., 802.
——ahena, sp. nu. (Pl. II. fig. 26),
802.
—— elevata, sp. n. (PI. II. fig.
802.
— neglecta, sp. n. (PI. II. fig. £
803.
Mimosella bigeminata,
(Pl. ILI. figs. 1-3), 851.
Mirounga leonina: facial vibrissx, 901.
Modiolus philippinarum, 666, 667.
Mo .uuscea :
Monte Bello Islands: systematic:
geographical, 665, 667.
Ostrea: ethology, 1061.
Zonitidee: Central Africa: systematic,
737.
Monitor gouldii, 642,
sje ie
INDEX.
Moncecocestus erethizontis, gen.
et sp. n.: structure (Figs. 1-9), 1039.
Mormosaurus seeleyi, gen. et sp.
n.: structure (Pl. TV. Figs. 1-4), 750,
767.
Murena thyrsoidea, 650. :
Mus musculus : facial yibrisse, 901.
rattus: ethology, 631.
avellanarius :
Muscardinus facial vi-
brissee, 902.
Myosoma spinosa, 804.
Myrrapopa :
Monte Bello Islands:
649.
systematic,
Nacaduba biocellata, 644.
Nakuruella soror, gen. et sp. n.
(Pl. I. fig. 13), 802.
Nanotragus pygmeus: facial vibrissz,
911.
Nataiina permembranaea, 800.
Natica vitellus, 666, 674.
Nawxia serpulifera, 661.
Naxioides serpulifera (P1. 11. figs. 9, 10),
development, 661.
Nectarinia famosa (z. t. s.), 1058.
Neosleptria punctifera, 646.
Neolucia serpentata, 644.
Nerita albicilla, 666.
Nettopus coromandelianus (z 1. s.), 943,
1057.
Nomia flaviviridis doddi, 649.
Notophoyx flavivostris (z.s.u.), 1059.
Nototragus melanotis: facial vibrissz,
SILL.
Nyctipithecus trivirgatus: facial
brisse (Fig. 6), 898.
vi-
Octodon degus: facial vibrissze, 903.-
Odatria ocellata, 642.
Opsanus diemensis, 650.
Orectolobus tentaculatus, 650.
Oreocincla lunulata (z.s. u.), 1058.
Ornithorhynchus: facial vibrisse, 891.
Orthederides, sp., 648.
Orycteropus capensis (2. s. L.), 1058.
| Oryetolagus cuniculus: facial vibrissee,
905,
INDEX.
Oryx gazelle (z.s t.), 1057.
Osmotreron griseicauda (z.s.x.), 1059.
* Ostreca cucullata, 656.
Ostrea: ethology, 1061.
Otaria californiana:
901.
Otiditenia eupodotidis: structure (Fig.
11), 879.
Otocyon megalotis: feet: rhinarium :
facial vibrisse (Fig. 8), 927.
Oxybelis acuminatus, 816.
brevirostris, 816.
Oxyrhopus petolarius, 816.
facial vibrissx,
Oxytoxia argenteo-ornatus, 645.
Pachycephala rufiventris collett?, 638...
Pagurus punctulatus, 656.
Pandesma submurina, 646. -
Pandion haliaétus melvillensis (Pl. 11.),
ethology,.639.
Paphia literata, 666.
Paracolletes perfasciatus, 648.
Paradisea apoda (z. 8. u.), 1059.
—— minor (z.s..1.), 1059.
Paralastor: ethology, 563.
abnormis,. 622.
albifrons, 566,584.
albocinctus, 586.
—— alexandrie, sp. n. (Pl. I. figs.
7, 22), 578, 618.
—— anostreptus, sp. n., 577, 613.
apicatus, 569, 584.
—— arenicolor, sp. n., 578, 618.
argentifrons, 573, 593.
-— argyrias, sp. n., 574. 597.
atripennis, sp. u. (PI. I. fig. 13),
574, 602.
aureocinctus, 574, 602.
australis, 566, 621.
—— bicarinatus, sp. n., 574, 601.
—— brisbanensis, sp. n., 575, 606.
—— brunneus, 577, 612.
—— carinatus, 576, 609.
— clotho, 566, 579.
—— cognatus, 569, 581.
—— commutatus, sp. n., 575, 608.
—— comptus, sp. n., 578, 617.
—— ——— rubescens, var. n., 578,
617.
XXL
_ Paralastor conspiciendus, sp. n.,
569, 581.
—— co7spicrus, sp.n., 569, 580.
—— constrictus, sp. n., 577, 615.
cruentatus, 566, 622.
—— darwinianus, sp. n., 577, 617.
debilis, sp. n. (Pl. IL. fig. 20),
573, 599.
debilitatus, sp. n., 576, 611.
dentiger, sp. n. (Pl. I. fig. 11),
574, 603.
despectus, sp. n., 572, 58).
—— donatus, sp. n., 572, 588.
—— dubiosus, sp. n., 569, 580.
—— dyscritias, sp. n., 577, 614.
—— elegans, sp. n., 569, 581.
—— emarginatus, 571, 585.
—— erturgus, 573, 592.
— euclidias, sp, n., 574, 598.
—— eugonias, sp. n., 574, 596.
—— eustomus, sp.n., 575, 604.
—— fallax, sp. n., 575, 606.
—— flaviceps, 566, 569, 583.
—— frater, sp. n., 571, 585.
JSraternus, 568, 53).
—- graeffei, 526, 622.
— habilis, sp. n., 569, 583.
—— hilaris, sp. n., (Pl. I. fig. 18), 574,
600. .
hirtensis, 567.
hirtiventris, (2>.
-_— icarioides, sp. n. (Pl. I. fig. 2),
578, 620.
—— ignotus, sp. n., 578, 621.
——imitator, sp. n. (Pl. I. fig. 10),
573, 578, 595.
infernalis, 568, 579.
—— infimus, sp. n., 575, 603.
insularis. 570, 585.
— lachesis, 566, 599.
—— leetus, sp. n., 571, 585.
— lateritius, 566, 576.
——~ leptias, sp. n., 578, 620.
—— mackayensis, sp. n., 575, 607.
—— maculiventris, 574, 601.
—— medius, sp. n., 575, 604.
— mesochlorus, sp. n., 577, 616.
—— mesochloroides, st. n.,
577, 616,
XX1V
Paralastor microgonias, sj. n.,
574, 596.
mimus, sp. n., 573, 578, 594.
—— multicolor, sp. n. (Pl. I. figs. 9,
19), 577, 612.
—-— mutabilis, sp. n., (PJ. I. fig. 8),
576, 610.
nautarum, 566, 573, 600.
—— occidentalis, sp n., 574, 598.
—— odyneripennis, sp. n., 576, 610.
—- odyneroides, sp. n., 576, 610.
Be olorissspn|( Gisele dtigs.4, 10)
575, 608.
optabilis, sp. n., (Pl. I. fig. 12),
574, 587.
—— ordinarius, sp. n., 571, 586.
orientalis, sp. n. (Pl. I. fig. 21),
574, 599.
—— pallidus, sp. n., 570, 584.
parca (Pl. I. fig. 1), 571, 585.
picteti, 577, 615.
—— placens, sp. n., 573, 592.
—— plebeius, sp. n., 576, 611.
—— princeps, sp. n., 575, 607.
pseudochromus, sp. n. (Pl. I.
fic. 14), 575, 605.
punctwlatus (Pl. I. fig. 17), 571,
586.
pusillus, 588.
—~—— roseotinctus, sp. n., 972, 591,
—— rufipes, sp. n., 568, 579.
sanguineus, 597.
—— saussurei, sp. n., 368, 579.
similis, 586.
— simillimus, sp. n., 578, 619.
—— simplex, sp. n., 573, 594.
—— simulator, sp. n., 572, 588.
smithi, 567, 622.
solitarius, sp. n. (Pl. I. fig. 6),
574, 600.
subhabilis, sp. n., 569, 583.
—— subobscurus, sp. n., 575, 593.
—— suboloris, sp. n., 575, 608.
——subplebeius, sp. n. (Pl. I.
fig. 15), 576, 611.
—— summus, sp. n., 975, 604.
—— synchromus, sp. n., 578, 619.
— tasmaniensis, 569, 582.
INDEX.
Paralastor tricarinulatus, sp. n.,
569,582.
tricolor, sp. nu. (Pl. I. fig. 5),
572, 590.
—— tuberculatus (PJ. I. fig. 3), 576,
585.
unifasciatus, 569, 581.
—— viduus, sp. n., 576, 609.
—— vulueratus, 575, 605.
—— vulpinis, 572, 587.
—— -—. excisus, st. n., 572, 587.
xanthochromus, sp. n., 077,
614.
xerophilus, sp. n., 572, 591.
Parthenope (Rhinolambrus) pelagica,
665.
Pedicellina gracilis, 855.
spinosa (Pl. IV. figs. 10, 11),
854.
Pelecanus conspicillatus, 638.
Percivalia mnyiroénsis, gen. et.
sp. n. (Pl. I. fig. 12), 806.
Periclimenes hermitensis, sp. n.
(Pl. I. figs. 1-3), 655.
Peripia variegata, 640.
Periplaneta concolor, 648.
Perodicticus potto:
898.
Petreeca leggei (z. s.u.), 1058.
pheenicea (z.s. u.), 1058.
Petrogale penicillata: facial vibrissie,
893.
Petrophila
1058.
Phalaropus hyperboreus (z.s. u.), 1061.
Phaneropis muelleri, 641.
facial vibrisse,
cinclorbyncha (4. 8. L.),
Phascolarctus cinereus: dental varia-
tion, 1U71.
Philander
891.
Phoca vitulina: facial vibrissx, 901.
Janiger; facial vibrisse,
Phocena communis: development,
1061.
Phocosaurus megischion: structure
(Figs 11, 18), 762.
Phorocanthia (?) senio, 647.
Phrygilus gayi (z. s. u.), 1060.
Phrynonax pecitonotus, 815,
INDEX.
Phyllochetopterus, 990.
——anglica, sp. n.: structure (Pl.
VIL, Figs. 9, 10, 12), 984.
— dioti: structure (Fig. 13), 986.
pictus: development: structure
(Fig. 13), 986.
prolifica, sp. n.: structure: de-
velopment: ethology (Pls. IV., V-
Fig. 11), 972, 980.
Phymodius ungulatus, 659.
Physignathus gilberti (Pl. I. figs. 4-7):
ethology, 641.
Pilumnus cerulescens, var., 660.
vespertilio, 660.
Pionus maximiliana (z. s. L.), 948,
1057.
Pisa serpulifera, 661.
Piscss:
Monte Bello Islands: systematic,
650.
Clupea harengus : abnormal, 1060.
Pisobia minuta ruficollis, 638.
Pitta nove-guines (z.s.L.), 1059.
Platypodia granulosa, 658.
Plotosus angwillaris, 650.
Pnigalion oweni, gen. et sp. n.:
structure (Figs. 5-7, 12, 16), 750,
767.
Podiceps cristatus :
(Ply Its, JE), aie
Polychrus spurrelli, sp. n., (PI. I.
fig. 3), 814.
Polydesma lawsont, 647.
marmarinopa, 647.
Pelypus sp., 680.
Pouyzoa :
Cyclostomata, Ctenostomata, Endo-
procta: B. EK. Africa : systematic :
structure, 831.
Polyzosteria sp., 648.
Potamogale velox : facial vibrissx, 895.
Precis vellida, 644.
Proboscidea : facial vibriss, 906.
Proboscina sertularoides, 839.
Procavia capensis :
(Fig. 11), 906.
Procolophon trigoniceps: structure (Pls.
I-III. ; Figs. 1-5), 735.
Propatria mundoides, 646.
courtship-habits
facial vibrissz
XXV
Protosquilla trispinosa (P). IT. figs. 11,
12), 663.
Protozoa :
Foraminifera from Kerimba Archi-
pelago (see ‘ Transactions’).
: ethology, 1069.
Proverruca vinculum, gen. et
sp. n. (Pl. I. figs. 1-9: Fig. 1), 946.
Pseudalopex gracilis?: feet: rhina-
rium: facial vibrisse (Fig. 6),
922:
Pseudochirus peregrinus: facial vi-
brissee, 893.
Pseudochromis fuscus, 650,
Pseudozethus, gen. n., 622.
australensis, sp. n., 623.
Pteromys sp.: facial vibrisse, 903.
Pteropus medius: facial vibrissz (Fig.
5), 897.
Ptilorhis intercedens: ethology, 1070.
Pyrrbula erythrocephala (z. s. u.), 943.
Rangifer tarandus (z. s. u.), 943.
Ratufa indica: facial vibrisss (Fig. 8),
903.
Reithrodon typicus: facial vibrisse,
902.
REPTILIA:
Choco, Colombia : systematic, 813.
Monte Bello Islands: systematic :
ethology, 640.
South Africa (fossil) :
systematic, 995.
Carnivorous Therapsids: systematic :
structure, 1021.
Chelonia :
structure :
structure: systematic,
1011.
Deinocephalia: systematic: struc-
ture, 749.
Lacerta czsaris from Canary Islands,
681.
Procolophon trigoniceps : systematic:
structure, 735.
Youngina capensis (fossil): syste-
matic: structure, 1072.
Rhabdometra cylindrica, sp. n.:
structure (Figs. 1-10), 859,
Rhadinea decorata, 816.
Rhamphastos cuvieri (z. s. u.), 943.
Proc. Zoou. Soc.—1914, No. LX XITI. 73
XXV1
Rhinobatus armatus, 650.
Rhinobothryum lentiginosum, 816.
Rhinocerotidx : facial vibrissa, 908.
Rhinopoma microphyllum: facial vi-
brissx (Fig. 5), 898; (z. s. u.), 943.
Ihodona bipes, 641.
Rhodostoma auris-felis, 667, 675.
thombocephalus letus, 649.
morsitans, 649.
Rhophalodon, 774.
Rhynchium australense, sp. u.,
623.
Salwis tuberculatus, 649.
Saragus sp., 647. |
Sarcophilus harrisi:
(Fig. 1), 891. |
Scalpellum vimineum, sp. n. (Pl.
I. figs, 10-12), 952. |
Scaphella hedleyi, nom. n., 674.
reticulata, 674.
——- volva, 667.
zebra, var., 667, 674.
Schizophrys dama, 662.
Sciurus prevosti: facial vibrisse, 903.
vulgaris: facial vibrissz, 903.
Scolopendra leta, 649.
morsitans, 649.
Scymnognathus whaitsi ?:
(Figs. 3-5), 1027.
Scymnosuchus whaitsi, 1027. |
Sepiadarium, 677.
auritum, sp. n. (Fig. 1), 678.
Sepioloidea, 677.
Serialaria semiconvoluta, 847,
Sertularia pustulosa, 851.
Sesamodon : structure (skull), 1025.
Sitta frontalis (z. s. u.), 1057,
Sorex araneus: facial vibrissx, 896.
Spelerpes parvipes, 813.
Speothos venaticus: feet (Fig. 1), 914.
Sphex australis, 649,
Spilotes megalolepis, 815.
pullatus, 815.
Spiochetopterus, 990,
Spreo hildebrandti (z. s, L.), LOGO.
Stigmatops indistineta perpleaa, 635,
Stomatopora: structure (PI. TV. fig. 7),
858.
facial vibrissx
structure
INDEX.
Strepsiceros imberbis (z. s. u.), 1058.
STRUCTURE:
Mammaria: Facial vibrisse, 889;
Canide: Ursids: feet: rhinaria:
facial vibrisse, 913; Martes
martes: M. foina, 1062; ‘Tri-
chosurus : Phaseolarctus: Chry-
sochloris : dental variations, 1071.
Reprinta: (fossil) 8. Africa, 995,
1011, 1021; Procolophon trigoni-
ceps, 735; Deinocephalia, 749;
Youngina capensis (fossil), 1072.
Pisces : Clupea harengus : abnormal,
1060.
Insncra: Lygus pabulinus: suction-
mechanism, 685.
Crustacra : Cirripedia, 950.
Motuusca: Cephalopoda, 677.
Potyzoa: ovicells, ete., 831.
Vermipea : Chetopteride, 955 ; Mo-
neecocestus, 1059; Rhabdometra,
Otiditenia, 859.
Subularia montebelloensis,
sp. n. (Fig. 1), 673.
Sylochelidon strenwus, 636.
Sylvilagus superciliaris : facial vibrissee
(Fig. 10), 905.
Sympetes sp. 647.
Symphyletes sp., 647.
Tamandua tetradactyla: facial vibrissz,
895.
Tapinocephalus atherstonei: structure
(Fig. 10), 762.
Tapirus indicus: facial vibrisse (Fig.
11), 906.
Tayassu (Dicotyles) tajacu: facial vi-
brissee (Fig. 12), 908.
Telepsavus sp.: structure (Figs. 6-8),
969.
Tervia folini, 844.
wrregularis (Pl. LV. fig. 8), 843.
Thalamita dispar, nom. n. (Pl. I.
fig. 4), 657.
savignyt, 657.
Thalarctos maritimus:
930.
Thecadactylus rapicauda, 814.
feet (Hig. 9),
INDEX.
Vhylacinus cynocephalus: facial vi- |
brissxe (Hig. 2), 891.
Thynnus sp. 649.
Tiliqua australis, 642.
Titanosuchus: structure (Figs. 17, 18),
769, 774.
Ferox (Pl. V.), 769.
Trachycystis fusco-olivacea, 803.
nigrotincta, 803.
Trachysphex (?) pilosulus, 649.
Tragulus kanchil: facial vibrisse (Mig.
13), 908.
Tremarctos ornatus: feet, 936.
thibetanus: feet:
(Figs. 11, 13), 932, 937.
Trichosurus vulpecula: cytology, 943;
dental variation, 1071.
fuliginosus: facial vibrissee
(Fig. 3), 893.
Tridacna elongata, 666.
Trochozonites buhambaensis,
sp. n. (PL. II. fig. 23), 804.
expatriata, sp. n. (Pl. II.
fig. 21), 805.
—-kempi, sp. n. (Pl. IT. fig. 22),
805.
— suturalis, sp. n. (PI. II. fig. 24),
805.
Tropidonotus fuliginoides (z. s. L.),
1059.
Tropidorhynchus corniculatus (z. s. L.),
1058.
Trox crotcht, 647.
Trynga ruficotlis, 638.
Turbo squamosus, 666, 669.
Typhlops ammodytes, sp. n. (Pl.
I. figs. 8-10), 642.
rhinarium
Uca forcipata (Pl. II. fig. 8), 661.
Urguessella, gen. n., 803.
—— capillata, sp.n. (PI. IIT. fig. 28),
S04.
— cuticularis, sp. n.
fig. 27), 804.
——- esau, sp. un. (Pl. III. fig.
804.
urguessensis, sp. n. (Pl. ILI.
fig. 25), 803.
(Pl. III.
XXVI11
Uromys bruijnii: facial vibrisse (Fig. 8)
901.
Ursus americanus: feet:
(Figs. 10, 13), 950, 937.
arctos: feet (Fig. 9), 935.
horribilis: feet, 935.
Utetheisa pulchelloides, 645. °
rhinariura
Valenciennea longipimnis, 650.
Valkeria uva, 853.
—— vidovici, 848.
Tallentinia now Huvallentinia, 944.
Vanessa kershawi, 644.
Varanus acanthurus, 642.
giganteus (z.s.u.), 1058.
gouldii: ethology, 642.
VARIATION:
Mamata: Gazella scemmeringi,
944.
VERMIDEA:
Cestoda : Moncecocestus: structure:
systematic, 1039; Otiditsenia :
structure : systematic, 879; Rhab-
dometra :
359.
Polychzta from N.E. Pacific: syste-
matic: development (Pls. I—-VL.:
Figs. 1-13), 955.
Verrucide : stiology, 950.
Vesicularia dichotoma, 848.
Voluta ohlita, 667.
Vulpes bengalensis: feet: rhinarium -
facial vibrisse (Fig. 8), 926.
vulpes: feet (Fig. 8), 925.
structure: systematic,
Xanthias atromanus, 659.
Xanthodes atromanus, 659.
Xenodon colubrinus, 816.
Xenorhynchus asiaticus (z. s. L.), 1058.
Youngina capensis, gen. et sp. n-
(Figs. 1, 2), 1075.
Younginide, fam. nov., 1077.
Zingis aurea, 802.
bullata, 802.
XXV1H
Zingis consanguinea, 796.
gaziensis, 795.
gregorti, 802.
kempi, 796.
papyracea, 799.
planispira, 800.
INDEX.
Zonitide: Central Africa (Pls. I.-III.),
787.
Zoobotryon pellucidum: structure (Pls.
II]. figs. 4-12; IV. fig. 12), 849.
Zostcrops balstoni, 635.
lutea balstoni, 635..
Zoneginthus castanotis roebucki, 636.
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LIST OF CONTENTS.
1914, Part III. (pp. 491-944). -
EXHIBITIONS AND NOTICES.
Page
The Srcrurary. Report on the Additions to the Society’s Menagerie during the month
Of May 19IA: eae seis oo ie inielernne tpiate cen elete = ie tedare as Rise teas wield ite 3s eee 943
Mr. J. Tuornron Carter, F.Z.8. Exhibition of Microphotographs of Mitosis in the Cells
of the Enamel Organ in Dasyurus viverrinus and Trichosurus vulpecula ......++.00- 943
Mr. R. 1. Pocock, F.R.S., F.L.S., F.Z.8., Curator of Mammals. Exhibition of a white
specimen of Sommering’s Gazelle ....+... 1.0... sere ees de alse, Co seine eae 944
The Rev. T. RB. BR. Stassine, M.A., F.R.S. Huvallentinia, nom. n. for Vallentinia Stebbing 944
PAPERS.
33. The Courtship-habits of the Great Crested Grebe (Podiceps cristatus); with an
addition to the Theory of Sexual Selection. By Jutian S. Huxury, B.A., Professor
of Biology inthe Rice Institute, Houston, Texas. (Plates I. & IL.)...... ote 491
34. On the Species of Alastor (Paralastor) Sauss.,and some other Hymenoptera of the
Family Eumenide. By R. C. L. Peruins, M.A., D.Sc., F.Z.8. (Plate I.) ...... e- 563
35. A Report on the Fauna of the Monte Bello Islands. By P. D. Monraeus, B.A.,
Gonville and Caius College, Cambridge. (Plates I-IV.) .........ceccssceececs 625 »
36. Stalk-eyed Crustaceans collected at the Monte Bello Islands. By Mary J. Rarusun,
United States National Museum, Washington, D.C., U.S.A. (Plates I. & II.) ...... 653
37. Report on Mollusca collected at the Monte Bello Islands, By Tom Irepaut, -
(Text-figure 1.) ....... PRR RE TERETE IM aT HE ORL LO GRECO EERE ee) oan 665 —
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COURTING-HABITS ‘OF PODICHPS CRISTATUS.
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33. The Courtship-habits* of the Great Crested Grebe
(Podiceps cristatus); with an addition to the Theory
of Sexual Selection. By Junian 8. Huxtey, B.A.,
Professor of Biology in the Rice Institute, Houston,
Texas f.
[Received March 14, 1914: Read April 21, 1914. ]
(Plates I. & IL.)
Inp5Ex.
J. GENERAL Parr. Page
APR tr ochuctrOne es ecceresctessees cas eter ca varscedecaceccvn cuts e sad cone AOU
Dor AN peaiian Cele vat My sesnuey setae acces eine ees aaeNeaecumcemcesverscs 402
3. Annual History .. Fas OS SON ees Re anish GEE SCaE Aine TOME meram oa)
4. Some Detouatane. Beeuaiannwenanec enact sstntescetes oes cees S200
6. The Relations of We exces
(Go) eihevactiotepanuinopeee ss eee eceee eet neces OOD
(Gis) Courtship iene see te sence eerie: Se nosatpeeeadccesweceet OOS
Gus) Nest=bunl din cesarean error teeecntesestastasiensea Oil/Z!
(iv.) Relations of different pairs .. ...........c00 eee SLL
(@eaynOthewmachivitiespaen cae tee tec e aes eeceweseienien MOLE
6. Discussion 522
* Tt was not until this paper was in print that I realized that the word Courtship
is perhaps misleading as applied to the incidents here recorded. While Courtship
should, strictly speaking, denote only axte-nuptial behaviour, it may readily be
extended to include any behaviour by which an organism of one sex seeks to “ win
over” one of the opposite sex. It will be seen that the behaviour of the Grebe
cannot be included under this. ‘‘ Love-habits”? would be a better term in some
ways; for the present, however, it is sufficient to point out the inadequacy of the
present biological terminology.
i Communicated by the SECRETARY.
{ For explanation of the Plates see p. 561.
Proc. Zoou. Soc.—1914, No. XXXV, 35
499, MR. J. 8S. HUXLEY ON THE
II. Sprcrar Parr. Page
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1. InrrRopucTION.
In these days the camera almost monopolizes the time and
attention of those who take an interest in the life of birds.
It has rendered splendid service, but I believe that it has almost
exhausted its first field. At the present moment both zoology and
photography would profit 1f naturalists for a little time would
drop the camera in favour of the field-glass and the note-book.
For the many who do not care alone using a telescope, the
prismatic binocular has more than doubled the possibilities of
field-observation ; and when full advantage shall have been taken
of those possibilities, not only will science be the richer for z
multitude of facts, but then, and only then, will the photo-
grapher, now hard-pressed for new subjects, suddenly find a
number of fresh avenues opened up to him.
This second paper on the courtship-habits of British birds,
like the first, will, I hope, help to show what wealth of interesting
things still lie hidden in and about the breeding places of familiar
birds. A good glass, a note-book, some patience, and a spare
fortnight in the spring—with these I not only managed to
discover many unknown facts about the Crested Grebe, but also
had one of the pleasantest of holidays. ‘Go thou and do
likewise.”
I shall first give a connected account of my own and others’
observations, followed by a discussion ; and in a second part or
appendix I shall give in detail some of the material worked
up in the first part, as well as some notes on various points not
connected with the main subject of the paper.
2. APPEARANCE,
Structure first, function afterwards: I must describe something
of the bird’s appearance before attempting to give an account of
its habits, though I shall try to be as brief as possible, since any
COURTSHIP OF THE GREAT CRESTED GREBE. 493
standard work of descriptive ornithology will give full details of
the plumage and taxonomic characters. The Great Crested
Grebe, then, is of course a water-bird, and essentially a diving-bird.
Its tail is remarkable in being reduced to a few tiny feathers,
and its legs are set as far back as possible, so as to have the
position of a ship’s propellers. Its body is long and approaches
the cylindrical; the neck is very long and flexible, the head flat,
the beak sharp, long, and powerful. In colour, the Great
Crested Grebe has back and flanks of much the same smoky
mottled brown as its small cousin, the Dabchick; the underparts,
however, including the chin, throat, and front of the neck, are of
an exquisitely pure white (furnishing the ‘‘ Grebe” of commerce).
The back of the neck is very dark brown.
The chief ornament of the bird, the crest from which it takes
its name, is reserved for the head. In these pages I shall use
the word crest to denote all the erectile feathers of the head taken
together. The crest, as thus defined, consists of two parts—the
ear-tufts (or ears, a8 for brevity’s sake they may be called) and
the ruff.
Both are composed of special narrow, elongated feathers, stiff,
and formed of comparatively few barbs. Those constituting the
ears are black, all of about the same length, and spring in two
tufts from the top of the head, above the tympanum. The ruff
is bigger and more elaborate: it consists of a broad band of
feathers springing from the sides of the face and head, their free
ends pointing downwards and backwards on either side of the
neck. If we take the part of the head behind the eye, we find
that at first the feathers are of the ordinary length, then slightly
elongated (the beginning of the ruff), and then longer and longer
till we get to the hinder border of the ruff. Corresponding to
the increase of length there is a change in colour., The proximal
(upper) part of the ruff is white, then we get to vivid chestnut,
and this deepens gradually to glossy black (see any good picture
of the Crested Grebe).
Both ruff and ears are extremely erectile; and as the birds
make great play with them during all the actions of courtship,
the various positions into which they can be put must be
described.
Let us begin with the ears. These, when depressed or shut,
stretch straight out backwards, continuing the line of the flat
head’s crown. When shut forcibly, the feathers of which they
are made are close together and all parallel.
Often, however, they are not thus “at attention,” but ‘standing
at ease,” to use a military metaphor: then the tufts as a whole
point in the same direction, but their component feathers diverge
and bristle-out a bit, This seems to be the usual and most
restful condition.
Further, the tufts may be erected: and they may be erected in
two ways—either laterally ov vertically. When erected laterally,
they stick out horizontally at right angles to the head, so that
35%
494 MR. J. S. HUXLEY,ON THE
from the sides they can scarcely be seen, as they are’ end-on to
the eyes. When erected vertically, they seem, when viewed from
the side, to be sticking straight upwards ; but when they are
seen from in front, it is found that they diverge from each other
at a considerable angle (Pls. I. & II. figs. 4, 5, 11). Durmeg
erection, the individual feathers always diverge fanwise very
considerably. Thus there are four conditions of the ear-tufts
to be distinguished.
They may be:—(a) Depressed.
(1) Shut tight.
(2) At rest (relaxed).
(b) Hrected.
(3) Vertically.
(4) Laterally.
The ruff is more complex in its attitudes, as im its structure.
During depression it, too, may be either shut tight or lying easy.
When really shut, it bears from the side a curious resemblance to
the gill-covers of some eel-like fish : its rounded hinder border lies
along the side of the cylindrical neck, whose outlines its own
scarcely overlap, either dorsally or ventrally (fig. 1). When re-
laxed (at rest), this resemblance disappears, for the feathers all
diverge slightly, and the smooth appearance of the surface is
lost.
When the ruff is erected, the feathers composing it may be
made to diverge in a single plane only, the original (longitudinal-
vertical) plane of the “ gill-cover,” or they may diverge outwards
as well, making an angle with the side of the head (movement in
the transverse, as well as the sagittal plane). I do not think that
they are ever moved in the transverse plane alone. Asa result
of these movements, three chief forms can be taken on by the
ruff. First there is the curtain form, m which motion in the
vertical plane alone takes place, the ventral edge being brought
forward till it makes an acute angle with the line of the chm
(fig. 3), the two halves thus hanging like curtains on either side
of the head. Then there is the pear-shaped condition (figs. 4,
11), where there is a consideyable amount of forward and a
moderate amount of transverse motion. The ruff in this state
has its vertical height greater than its breadth (fig. 5). Owing
to the transverse bristling of the feathers, the two halves of the
vuff almost blend into a single whole; they can scarcely be
distinguished either from the front or, stall less, from behind,
whereas in the curtain form they are very distinct. Finally,
there is the elliptical form, when, added to the same amount of
Jongitudinal motion, the greatest possible amount of transverse
bristling has taken place. The ruff is now actually broader than
it is high (fig. 9), and the blending of the two halves is practically
complete. There are, of course, intermediate states. Instead
of “full pear-shaped,” you may have ‘half pear-shaped ” ;
Ly | ; and
between pear-shaped and elliptical there comes the circular.
COURTSHIP OF THE GREAT CRESTED GREBE. 495
The three I have named, however, are those which the bird
usually adopts.
The ruff, therefore, may be :—
(a) Depressed.
(1) Shut tight (‘ gill-cover”).
(2) At rest (relaxed),
(b) Hrected.
(3) Curtain-like (motion of feathers in one plane).
(4) Pear-shaped
(5) Elliptical
By a combination of particular positions of ruff, ears, and neck,
and sometimes wings and body too, the birds can assume a
number of characteristic and often-recurring attitudes, which are
the raw materials, so to speak, of all the elaborate habits of
courtship. j
Before giving any more definitions, I will now give an outline
of the Grebe’s annual history, and then go on to describe some of
the actual happenings that I saw, in order to give an idea of the
problems to be solved. Then I shall try to define and classify
the various courtship-habits, and discuss the general bearing of
the facts.
} (motion of feathers in two planes).
3. ANNUAL History.
This is somewhat as follows *. About the first week of
February they leave the sea-coast and fly back in bands to the in-
land waters where they breed. They live in flocks for about three
weeks, and then start pairing-up. Pairing-up lasts altogether
about a fortnight, bringing us to mid-March. From this time
ou to the end of summer, the unit is neither the flock nor the
individual, but the family, represented at first by the pair.
About the beginning of April nest-building begins, and by the
end of the month every nest will have eggs. The family parties
live together through the summer, though apparently the cock
leaves the hen to look after the young when they are half-grown
(Pycraft, ’11). There is usually no second brood unless the first
is destroyed. At the end of September they gather into flocks
again, and live thus for well over a month, finally leaving for the
sea-coast in the second week of November.
The period of pairing-up itself I have unfortunately not been
able to observe; the keeper tells me that there is much flying and
chasing about. The part played by the “courtship” in the actual
pairing-up is thus left uncertain. From analogy with other birds
and with ourselves we should expect that the chasing was the
expression of felt but unreasoned likes and dislikes, and that the
courtship-actions were only gone through after the two birds had
become fairly well-disposed towards each other. The courtship-
* The dates refer to the movements of the birds at Tring Reservoir, and have
been given me by the head-keeper there.
496 MR. J. 8. HUXLEY ON THE
actions, I am told, are at any rate to be seen immediately after
pairing- up.
4. Some DescrrPrions.
(a) Let us start with the commonest of all the scenes of court-
ship—the one which had first attracted and puzzled me years ago,
and led me to choose the Grebe as a bird to watch.
As the birds ride on the water, very little of their under-
surface is usually visible; but now and then a twinkle of white is
seen. This may be merely a bird rolling half over to preen its
belly; but if it proceed from two birds close together, this form
of courtship is almost sure to be in progress. In such a case, the
glass reveals that the two birds are always a pair, cock and hen ;
they are facing each other, their beaks perhaps a foot, perhaps a
mere couple of inches apart, their necks held up perfectly straight
and elongated to a truly surprising extent. It is this holding up
of the neck that shows some of the white of throat and breast.
Their ears ave erected vertically and their ruffs are full pear-
shaped. he few little feathers that do duty for a tail are
cocked up as far as they will go—that is to say, about half an
inch (fig. 11).
In this attitude the birds proceed to go through a curious set
ritual.
Let us describe a particular case. A pair of birds, cock and
hen, that had been fishing not far apart, suddenly approached
each other, raising their necks and rufts as they did so, till by
the time they had got face to face they were in the attitude I
have just described. Then they both began shaking their heads
at each other in a peculiar and formal-looking manner. Each
bird began by waggling its head violently from side to side,
some four or five times in quick succession, like a man nodding
emphatic dissent. Then the quick side-to-side motion gave place
to a slow one, and the beak and head were swung slowly across
and back, with a seemingly vague and enquiring action, as if the
bird were searching the horizon for it knew not what. The head
was moved back and forth perhaps a couple of times, and then
the violent shaking beganagain. This alternation of shaking and
slow side-to-side swinging was repeated over and over again by
each bird: strangely enough, the pair kept no time with each
ether—the violent shakings of the two neither coincided nor
alternated, but each shook and swung without any apparent
reference to the other’s rhythm.
After six or seven repetitions of the performance another
action came in. After the slow swing and before the wageling
(or sometimes, I think, taking the place of the slow swinging),
but not every time, the bird bent its neck right back and down as
if to preen its wings, put its beak under some of the wing-feathers
near the tail, raised them an inch or so, let them fall, and brought
its head swiftly hack into position for another of the violent
shakings. This action had obviously something to do with
COURTSHIP OF THE GREAT CRESTED GREBE. 497
preening, but had an extraordinary look, as of a stereotyped
and meaningless relic. The birds seemed to be performing some
routine-action absent-mindedly and by mere force of association,
as one may sometimes see a man wind up his watch in the day-
time, just because he has been changing his waistcoat.
Finally, after each bird had given about a dozen or fifteen
violent shakes, with a corresponding number of slow swings and
liftings of wings in between, they veered up into the wind almost
simultaneously, lowered their crests, brought their necks down,
and, in a word, became normal once more both in appearance and
behaviour.
One must have names for things if one is going to discuss them,
as any philosopher will tell you. So I propose to call a whole
performance such as that just described a bout of shaking; each
little time of violent wagegling I shall call simply @ shake, and
shall measure the length of a bout by the number of shakes in it
(counting the shakes of both birds added together). Finally, the
curious actions resembling preening I shall call habit-preening,
because I believe them to have (or rather, to have had) something
to do with real preening, but, in the per formance as gone through
to-day, to have "eCoUne & : mere habit, vestigial so far as its or iginal
function is concerned.
(5) That is one little seene: now take another.
A solitary bird, which proved to be a hen, came flying over from
one reservoir to another. She alighted near one shore and began
swimming slowly across towards the other, meanwhile alternating
between two attitudes.
First of all she arched her neck right forward till the bill,
which pointed slightly downwards, was just above the water.
The ears meanwhile were scarcely erected ; the ruff was thrown
forward in curtain-form, and thus, since the head had been
brought so far forwards and downwards, actually swept the
water on either side (fig. 3). So she progressed, looking from
side to side, and now and then giving a short barking call.
After four or five of these calls, which represented perhaps 20 or
30 seconds of time, she put down her ruff and raised her neck
nearly straight up to enlarge her circle of vision.
After some seconds of looking about her in this position, she
relapsed again into the first attitude—‘ with neck outstretched,
you fancy how.” This was repeated eight or nine times, til at
last a cock, thirty or forty yards away, appeared to notice the
ealling bird. He pricked up his neck, looked towards her for a
short “une. and then dived. At eh she changed her whole
demeanour. Up went her wings: back hebween them, with
erected ruff and ears, went her head. A glance at fig. 7 will
show her attitude. The wings were brought up, half- spread on
either side of the body, with their antorion border pointing down-
wards. They were almost in the transverse plane, but sloped
slightly backwards from the water.
498 MR. J. 8S. HUXLEY ON THE
In this position the beautiful white bar formed by the marginal
wing-coverts, along the anterior margin of the wing, and the
broad white blaze formed by the secondaries, which are quite
invisible when the wings are closed, shone out vividly. The gap
between the wings was filled by the head ; this from the front
somewhat resembled an old-fashioned picture of the sun, with the
ruff rayed out considerably all round, and the ears were erected
laterally so as to fit on to the top of the ruff on either side.
Below the head shone the white of the puffed-out breast. The
bird’s whole appearance was wonderfully striking, and as unlike
as possible to that of its everyday self.
All this took but an instant; directly the cock had dived she
was in this attitude. As she waited for his re-appearance she
turned eagerly from side to side, swinging nearly to the right-
about and back again as if not to miss him. Kventually he came
up, three or four feet on the far side, and facing away from her
in the most amazing attitude. I could scarcely believe my eyes.
He seemed to grow out of the water. First his head, the ruff
nearly circular, the beak pointing down along the neck in a stiff
and peculiar manner; then the neck, quite straight and vertical ;
then the body, straight and vertical too; until finally the whole
bird, save for a few inches, was standing erect in the water, and
reminding me of nothing so much as the hypnotized phantom of
a rather slender Penguin.
As I say, it grew out of the water, and as it grew it gradually
revolved on its long axis until at its fullest height it came to face
the hen. Though all this was done with an unhurried and
uniform motion, yet of course it took very little time. Then
from his stiff, erect position he sank slowly on to the surface; the
hen meanwhile put down her wings and raised her neck; and the
pair settled down to a bout of the head-shaking. Their attitudes
and actions were practically the same as those of the pair described
above, but the bout only lasted about half as long. It was ended
by the two birds ceasing to shake and gradually drifting apart.
Finally they put down their crests and went off together to preen
themselves and fish.
These actions, too, must now be named. I propose to call
the attitude of the hen as she searched and called for her mate
the Dundreary-attitude, for the two halves of the ruff in curtain-
form give the bird, especially when seen from the front, a con-
siderable resemblance to that famous personage of the drama.
The attitude later assumed by the hen, with head back and wings
arched, shall be the Cat-attitude, for the round ruff gives the bird
the look of a very contented and somewhat fat cat. The cock’s
combined dive and emergence I shall call the Ghost-dive. The whole
ceremony I have called the Discovery Ceremony (see p. 512).
(c) Now for the highest development of the courtship-actions
that I have seen. The incident I am going to describe took place
in the middle of the hour-and-a-half’s watching the results of
COURTSHIP OF THE GREAT CRESTED GREBE. 499
which are recorded on p. 549. A pair of birds had been preening
themselves and fishing, with occasional languid bouts of head-
shaking. After a dive they came up not far apart and swam to-
gether with outstretched necks, which, as they neared each other,
they gradually raised, beginning to shake their heads a little at
the same time. The raising and the shaking progressed simul-
taneously, till when the birds were face to face they were in the
regular ‘“‘shaking” attitude and waggling their heads with a
vengeance. The bout of shaking thus begun was the longest I ever
saw : between them the birds shook their heads no less than 84
times, and with as much vigour at the eighty-fourth as at the first
shake. There was rather a curious difference between the cock
and the hen. At first neither of the birds did any of the wing-
lifting, the strange parody of preening described above, and
there named habit-preening. After the fifteenth shake, the hen
began to give an occasional wing-lift, and these became more and
more frequent on her part, until after about the sixtieth shake
she was turning round and putting her beak under her wing-
feathers between nearly every shake. The cock, on the other hand,
did not begin this habit-preening until after the fortieth shake,
and even after that only repeated the trick at rare intervals.
At the close of the bout, the pair swung parallel, but did not
bring their necks down. Nor did they lower their ruffs; on the
contrary, they put them up still further, from the pear-shaped
form customary for shaking to the extreme elliptical, bringing
down their ears meanwhile from the vertical to the lateral posi-
tion, so that the whole crest now appeared like a large chestnut-
and-black Elizabethan ruff. This change in the crests made me
think something exciting was going to happen. Sure enough,
the hen soon dived. The cock waited in the same attitude,
motionless, for perhaps a quarter of a minute. Then he, too,
dived. Another quarter of a minute passed. Then the hen
appeared again, and a second or two later, some twenty-five yards
away, the cock came up as well.
They were in a crouching position, with necks bent forward,
ruffs still elliptical, and both were holding in their beaks a bunch
of dark ribbony weed, which they must have pulled from the
bottom. The hen looked about her eagerly when she first came
up: when the cock appeared she put her head down still further
and swam straight towards him at a good pace. He caught sight
of her almost immediately too, and likewise lowering his head,
made off to meet her. They did not slacken speed at all, and I
wondered what would happen when they met. My wonder was
justified : when about a yard apart they both sprang up from the
water into an almost erect position, looking somewhat like the
‘“‘shostly Penguin” already described. Sprang is perhaps too
strong a word; there was no actual leap, but a very quick
rising-up of the birds. The whole process, however, was much
quicker and more vigorous than the slow “ growing out of the
water ” of the ghost-dive. In addition, the head was here not
500 MR. J. S. HUXLEY ON THE
bent down along the neck, but held slightly back, the beak hori-
zontal, still holding the weed. Carrying on with the impetus of
their motion, the two birds came actually to touch each other
with their breasts. From the common fulcrum thus formed
bodies and necks alike sloped slightly back—the birds would have
fallen forwards had each not thus supported the other. Only the
very tip of the body was in the water, and there I could see a
great splashing, showing that the legs were hard at work. The
appearance either bird presented to its mate had changed alto-
gether in an instant of time. Before, they had been black and
dark mottled brown : they saw each other now all brilliant white,
with chestnut and black surrounding the face in a circle.
In this position they stayed for a few seconds rocking gently
from side to side upon the point of their breasts; it was an
ecstatic motion, as if they were swaying to the music of a dance.
Then, still rocking and still in contact, they settled very gradually
down on to the surface of the water; so gradually did they sink
that I should think their legs must have been continuously
working against their weight. All this time, too, they had been
shaking their heads violently at frequent intervals, and after
coming down from the erect attitude they ended the performance
by what was simply an ordinary bout of rather excited shaking ;
the only unusual thing about it was that the birds at the begin-
ning were still, I think, actually touching each other. The weed
by this time had all disappeared: what had happened to it was
very hard to make out, but I believe that some of it was thrown
away, and some of it eaten by the birds while settling down from
the Penguin position.
In their final bout they shook about twenty times, getting less
excited towards the end ; they eventually drifted apart, put their
crests down, and almost at once began to pick food off the surface
of the water.
Let us call the diving for water-weed and the appearing again
with it in the bill the weed-trick ; and the rapid swimming to-
gether, with the subsequent figure erect breast-to-breast, let us
call the Penguin-dance, for here once more the general resemblance
to Penguins (exceptionally graceful ones, let us admit) forced
itself upon the mind.
(d) One last scene before we pass from mere description to the
heavier task of analysis.
Sitting on the bank one day, looking ‘out over a broad belt of
low flags and rushes which here took the place of the usual
Arundo, I saw a Grebe come swimming steadily along parallel to
the bank, bending its head forward a little with each stroke, as is
the bird’s way in all but very leisurely swimming. I happened
to look further on in the direction in which it was going, and
there, twenty or thirty yards ahead of it, I saw what I took to be
a dead Grebe floating on the water, The body was rather humped
up; the neck was extended perfectly straight in the line of the
COURTSHIP OF THE GREAT CRESTED GREBE. 501
body, flat upon the surface of the water; the ruff and ears were
depressed (fig. 8). So convinced was I that this was a dead bird
that I at once began revolving plans for wading in and fetching
it out directly the other bird should have passed it by. Mean-
while, I wanted to see whether the living would show ary interest in
the dead, and was therefore much interested to see the swimming
bird swim up to the tail-end of the corpse and then a little way
alongside of it, bending its head down a bit as if to examine the
body. Then it came back to the tail-end, and then, to my
extreme bewilderment, proceeded to scramble out of the water on
to the said tail-end; there it stood for some seconds, in the
customary and very ungraceful out-of-water attitude—the body
nearly upright, leaning slightly forward, the neck arched back and
down, with a snaky Cormorant-look about it, the ruff and ears
depressed. Then it proceeded to waddle awkwardly along the
body to the head end, slipping off thence into the water and
eracefulness once more. Hardly had it done this when the
supposed corpse lifted its head and neck, gave a sort of jump, and
it, 400,.was Swimming in the water by the other’s side. It was
now seen that the “ corpse” had been resting its body on a half-
made nest whose top was scarcely above the water, and it was
this which had given it the curious hunched-up look. The two
swam about together for a bit, but soon parted company without
evincing any further particular interest in each other.
Both these birds had crests of very much the average size, so
that it was hard to tell their sex ; but I think that the “ corpse”
was a hen, the other bird a cock.
The meaning of this action (which I only saw this one time)
remained extremely problematical to me while I was at the
Reservoirs. The mystery will, however, be solved in the next
section, and so let us anticipate and call the attitude of the
“corpse” the passive, that of the bird that climbed on the
eS)
*“ corpse’s ” back the active pairing attitude.
5. Tar RELATIONS OF THE SEXES IN THE GREAT CRESTED GREBE.
(i.) The Act of Pairing.
As I say, it was especially the proceeding last described which
puzzled me; and it was not till I had got home and looked up
the literature, that I found a welcome paper by Selous (01) *
which exactly dovetailed into my own observations. I had
been mainly concerned with the behaviour of the birds on
the open water and during incubation; he had paid special
attention to nest-building and pairing His observations solve
the mystery that has so far surrounded the Grebes’ actual
pairing ; by them it is now established that the attitude which
so puzzled me is adopted always, and only, for the purpose of
coition, and that coition takes place solely on the nest. 1 should
* A short summary of this paper will be found on p. 529,
502 MR. J. S. HUXLEY ON THE
perhaps have said “on a nest”; for the birds may build several
incomplete nests or platforms before one, finally chosen to be
the true nest, is finished and laid in.
From Selous’s observations, the actions and ceremonies con-
nected with coition are quite elaborate—almost of the same
order of elaboration as the courtship-ceremonies, though the
two rituals are completely independent and appear to have
developed along quite different lines.
We have already got to know the passive and the active
pairing attitudes. To complete the description of the mere
attitudes, it remains to add that, before sinking down into
the passive pairing attitude (which Selous calls “lying along
the water’), the birds usually assume a curious fixed and rigid
pose. I will quote Selous’s words:—“. . . curling his neck over
and down, with the bill pointing at the ground [weeds], perhaps
six inches above it, he stood thus, fixed and rigid, for some
moments (as though making a point) before sinking down and
lying all along. There was no mistaking the entirely sexual
character of this strange performance, the peculiar fixed rigidity
full of import and expression.”
We must now see how the attitudes are combined in the
actions themselves.
In the first place, we have the active pairing attitude and
the actions associated with it. These actions in pairing have
been already once described, and they seem to show little
variation. The bird leaps up, and comes down almost upright
near the other’s tail. Copulation is then attempted. Selous
found it hard to decide if such an attempt was successful or
not. When it seemed successful, the birds apparently uttered a
louder cry than usual, and afterwards their behaviour had a
satisfied look. Once, however, when the birds seemed thus
satisfied, he adds: ‘‘The time occupied was extremely short,
and one would hardly have thought from the position of the
two birds that actual pairing had been possible.” In other
cases he could be fairly sure that the attempt was not successful.
Whether successful or not, the act always ends in the peculiar
way already described: the active bird waddles forwards along
the other’s body, and walks somehow over its head into the
water, upon which the passive bird raises its neck, leaves the
nest or platform, and swims away in normal position.
In the second place, we find that the passive attitude (‘lying
along”) may take place either on the nest or platform itself, or
else on the open water (but then apparently never far from the
nest); the act of pairing itself, however, is possible only when
the *‘ passive” bird is lying on some firm support. In the second
place, both cock and hen go into this attitude (the precise attitude
which the lower bird assumes during the act of coition) indis-
criminately : Selous’s records give an approximately equal number
of times for the two sexes.
However, before trying to draw any general conclusions, let us
COURTSHIP OF THE GREAT CRESTED GREBE. 503
take a particular case—that described by Selous on pp. 180-181
of his paper :—
About an hour earlier in the day there had been an attempt at
pairing. Then, after a period of rest on the open water, the
birds swam together towards the nest (which had been built
the day before). When just outside the bed of reeds in which
the nest was situated, the hen went into the passive attitude,
on the open water. The cock came up to her, swam a few yards
past her, went twice back to her and away again, then went
right into the weeds and himself lay along the water in the
passive attitude. While he was doing this (or immediately
afterwards) the hen swam to the nest, leapt on to it, and sank
down in the passive attitude once more. Upon this the cock
came up to the nest, jumped on to the hen’s back, and they
apparently paired successfully, both birds meanwhile uttering a
special shrill screaming cry.
Here are various points to be noticed. The joint approach of
the birds to the neighbourhood of the nest is invariable when
they have previously been some distance away. When one bird
is sitting, or when both are already close to the nest, as when
building is in progress, the case is of course different (p. 533);
but in the period between nest-building and incubation they
seem never to approach the nest singly.
The passive attitude on the open water close to the weeds and
nest may or may not be assumed. In the three cases where
this happened and Selous is absolutely sure of his facts, the bird
that assumed this position was the female, and was also the
leader in the procession towards the weeds. (We want to know
more about this. It seems probable, from other considerations,
that it is a mere coincidence for the leader to have been always
the hen; but, this being granted, it is quite likely that the
leading bird would be the more eager, and so would hasten to
put itself into the attitude which apparently expresses readiness
to pair.) In other cases the birds swam straight to the nest,
and one of them ascended it and then went into the passive
position.
Next, the way in which the cock swam about close to the hen
while she was in the passive attitude, but still on the open water,
“as though about to pair” (I quote Selous), is interesting.
There must be a strong association established between the sight
of the passive attitude and the desire to pair, so that the active
bird shows its thoughts, so to speak, even when pairing is
impossible (as when the passive bird is on the open water).
When the passive bird has gone into position on the nest, it is
very nearly always the case that the active bird comes up to the
passive one and examines it or swims about a bit, whether an
attempt to pair is afterwards made (Selous, ’01, pp. 180, 345) or
not (loc. cit. pp. 165, 344, etc.). Sometimes the second bird is not
eager, and refuses to come near at all (¢. 9., loc. cit. pp. 172, 456).
At other times (loc. cit. p. 341, and perhaps p. 181) an attempt
5O4 MR. J. 8S. HUXLEY ON THE
to pair is made, and the active bird jumps up apparently
at once, without any delay. In the case observed by me the
active bird seemed very definitely to examine the passive one,
poking its beak down close to it; but the examination was very
short, the attempt at pairing following immediately.
When the active bird is moderately eager but not quite eager
enough to attempt to pair, it may swim up to the passive bird
a number of times, each time make as if to spring up, and then
decide not to, but swim away again.
The assumption of the passive attitude by one bird is generally,
so far as I can see, used as an invitation to the other bird to
pair: perhaps I should express myself rather differently, and say
that it always denotes readiness to pair, and is generally used as
a primary excitant—i. ¢., it is the first sign given by either of the
birds of readiness to pair. This is well brought out by incidents
such as this:—Both birds are building the nest; suddenly the
hen jumps up on to the nest and goes into the passive attitude,
every now and then raising her neck and looking round at the
cock (Selous, op. cit.). At other times it may be only a secondary
excitant—a mere symbol, This may happen when one bird is
sitting and the other approaches the nest; the sitting bird may
then assume the passive position at each approach of the other.
Here the approach is the primary stimulus, and the assumption
of the passive attitude is called forth by it, and not by internal
causes. In this second case a less degree of ‘‘ sexual feeling” is
presumably needed to induce the passive attitude than in the
first case.
But we are going too fast. We must not omit to notice the
curious action of the cock in himself copying the hen’s passive
attitude. This action—one bird going into the passive attitude,
the other coming up and examining it, and then going off and
assuming the same attitude—appears only to occur when the first
bird goes into position on the open water (and not on the nest),
and even then not always. It seems, however, to happen in the
majority of cases (though we are perforce generalizing from very
few instances). It looks as if it were a signal to the first bird
that the second was ready and willing to proceed further in the
matter; for the birds after this may proceed together to the nest,
where the first (Joc. cit. p. 180) or the second (ibid. p. 456) bird
ascends the nest and assumes the passive attitude once more.
In other cases, however (ibid. pp. 179, 454), the affair ended with
the second bird’s assumption of the attitude. Here it looks as if
a ritual ceremony was developing out of a useful action (see
below).
As regards the actual act of pairing (or its attempt, which for
our present purpose comes to the same thing), the two sexes seem
here also to play interchangeable roles. In 1900 Selous saw three
attempts to pair, one apparently successful, two unsuccessful :
in all three cases the active bird was the larger of the pair..
In 1901 he saw two attempts, both of which he thinks were
COURTSHIP OF THE GREAT CRESTED GREBE, 505
successful: here the active bird was in both cases the smaller of
the pair.
Now, if we could be sure that the 1901 pair was the same that
was there in 1900, all would be well: but we cannot be sure.
There was a marked difference in the pairing-behaviour of the
1900 and 1901 pairs—a difference that cannot be referred back
to the fact that in 1900 the birds were building a true nest and
were incubating, while in 1901 they had only got to the length
of building a pairing-platform. In 1900 the smaller bird (that
we have so far presumed to be the hen) was more forward in
invitation, while the active pairing-position was adopted by the
larger bird alone; im 1901 the case was exactly reversed. It
would be, in my opinion, more remarkable that such a change
of character should take place in two birds in the space of
one year than that the same water should be occupied by two
different pairs—albeit but a single one—in two successive years.
Mr. Selous, however, writes to me that for various reasons
(e. g., the site of the nest, etc.) he is practically convinced that
the birds were the same in both years. However, whether the
pair was the same pair or not, in both years there was a marked
difference in size both of body and crest between the two birds
of the pair, and, if all the books are not wrong, this should be
quite enough to distinguish the sexes. Sometimes, it is true,
the two birds of a pair are almost exactly alike; but nowhere
do I find it stated that the hen is ever larger or has a better
crest than the cock. It is the part of the professional orni-
thologist to find out if this is ever so; till then, we must be
content to say that it is extremely probable that either cock or
hen can play the “active” part in copulation—what we should
usually call the male part. This can be more easily imagined
in birds than in almost any other animals in which copulation
takes place, but even in a bird is remarkable enough. Definite
attitudes of the two participating organisms have been evolved to
facilitate the passage of genital products in a definite direction :
and here, hey presto! although the genital products continue to
pass in the same direction, yet the attitudes, developed only in
relation with and accessory to this direction, are at will reversed.
This facultative reversal of pairing-position would certainly be
remarkable; but even for the moment supposing that it does
not occur in our Grebe, it would merely appear as the as yet
unattained end of a process of sex-equalization which in this
species has already run a considerable course. This process
consists in a gradual transference of all the secondary sexual
characters of the male to the female, and vice versa. In its
general aspect it will be discussed later; here it will be sufficient
to consider it in relation to the pairing actions alone.
Let us see what is without doubt common to both sexes in the
Crested Grebe to-day. First of all, we find that either cock or
hen may lead the way towards the pairing-platform. Secondly,
either cock or hen may assume the passive pairing-positien (the
506 MR. J. S. HUXLEY ON THE
position that one would naturally call Jemale) on the open water.
Thirdly, either cock or hen may assume this position on the nest
or pairing-platform. This is important, for the pairing-platform
is never ascended except for the purpose of pairing, or for this
position, which we may call the beginning of, or the invitation to,
pairing, and the nest only ascended for these two purposes and for
incubation. Fourthly, when one bird is in the passive position,
the other, be it cock or hen, may come up to it, examine it, and
make as if to leap up on to it, just as it often does before an
actual attempt at pairing is made. The natural end of this
sequence would be that, fifthly, either cock or hen might not
only make as if to ascend into the active position, but actually
do so. If the text-books are right in their descriptions of the
sexes in the species, then we can say that this end has been
reached, and that, as far as pairing-positions go, the sexes are
interchangeable. If the text-books are wrong, then our evidence
is simply insufficient. Here it can only be shown that, however
incredible this reversal may appear, yet it is quite certain that
in the Great Crested Grebe all the preliminary steps towards
it have been already taken.
Further, Selous (loc. cit.) places on record some remarkable
facts which show that reversal of pairing-attitude does take place
in tame Pigeons. Here he several times saw, immediately after
the act of pairing, the “male” bird crouch, and the ‘“ female”
then get into the normal male attitude. The act of pairing was
then gone through a second time, but with the attitudes of the
birds reversed. See also Selous ’02.
We have therefore evidence that the full reversal can take
place, aud now only want to be certain that it has taken place in
this species. In any case we can say that characters (in this case
attitudes and actions only) of the female have been transferred to
the male, as well as characters of the male to the female.
We must now go on to consider a very different question,
which is also well brought out in the pairing-habits of the Great
Crested Grebe: I mean the gradual change of a useful action
into a symbol and then into a ritual: or, in other words, the
change by which the same act which first subserved a definite
purpose directly comes later to subserve it only indirectly
(symbolically), and then not at all. The action in question
here is the passive pairing-attitude, and the Grebe is interesting
as showing all three stages of the process at one time—the
passive attitude employed sometimes directly, sometimes sym-
bolically, and sometimes ritually. Speaking phylogenetically,
we have the following steps :—
(1) The ascent on to a nest or platform, and the assumption
of the passive attitude, are necessary if pairing is to take place,
and the passive bird must get into position before the active
bird can even begin its part in the coition act.
(2) The ascent and the attitude are used by the passive bird as
COURTSHIP OF THE GREAT CRESTED GREBE. 507
an incentive to the active bird, as a sign of readiness to pair.
The active bird may or may not respond.
(3) As a symbol, the attitude is obviously more important than
the actual ascent on to the nest, since the attitude is used only in
pairing, while the birds may ascend the nest for various purposes;
and, in addition, the assumption of the attitude comes after the
ascent, and is thus in time more immediately associated with
the act of pairing. Thus the attitude by itself comes to be used
on the open water (though always close to the nest) as a sign of
readiness to pair. We may say that readiness to pair is indicated
precociously —it is pushed back astep. Such processes of pushing
back are very common in early ontogeny ; embryologists then say
that the time of appearance of the character is cenogenetic (even
though the character itself, as here, may be palingenetic). The
phylogenetic change has here been precisely similar ; the only
difference is that the displacement affects a mature instead of a
very early period of lite.
(4) The attitude being now sometimes a mere symbol can be,
and is, employed by either the active or the passive bird. In
fact, when one bird employs it thus symbolically, the other
usually responds by immediately repeating this symbolic use.
(5) From useful symbolism to mere ritual is the last step—one
that has taken place often enough in various human affairs. It
appears that these actions and attitudes, once symbolic of certain
states of mind and leading up to certain definite ends, lose their
active symbolism and become ends in themselves. When I say
that they lose their active symbolism, I mean that they are now
not so much associated with readiness to pair as with the vague
idea of pairing in general. Thus associated with pleasurable and
exciting emotions, they may become the channels through which
these emotions can express themselves, and so change from
purposeful stimuli to further action into merely pleasurable self-
exhausting processes (see below). It is at least hard to see how
to explain such happenings as that described on p. 534, (c) 6,
where first one bird and then the other goes into the passive
position on the open water, after which there is simply a
resumption of feeding or preening.
Another general point worth noticing is this:—-In the case of
this Grebe the male has even less possibility of enforcing his
desires than the majority of birds. In a few birds the male is
not so helpless. The ordinary Barndoor Cock, for instance, is
often rather forcible in his methods. In the Wild Duck (Anas
boschas L.) the drakes often kill the ducks by continued tread-
ing*. Somewhat similar forcible pairing is recorded of the
Mute Swan (Cygnus olor). In such species it is by no means
necessary for the race that the act of pairing should be par-
ticularly pleasant to the female. In most birds, however, the
female has the upper hand: she can always prevent the cock
* Huxley, Biol. Centralbl. 1912.
Proc. Zoot. Soc.—1914, No. XXXVI. 36
508 MR. J. S. HUXLEY ON THE
from pairing with her, by simply running or flying away
(cf. the Redshank, Huxley, ’12,). In our Grebe we are a step
further still: not only must the female (or the passive bird, if we
want to be precise; but this is, for the present, complicating the
issue unnecessarily) be willing to pair, but she must also take the
first steps-—-must ascend a nest or platform and assume a special
position—before the cock can think of pairing. Here, therefore,
supposing that the functions of the sexes had not been almost
equally distributed, it would have been necessary for the hen to
have had a strong impulse towards pairing ; it might be that she
was impelled directly by a violent physiological stimulus, or
more indirectly by association, through the act being extremely
pleasurable.
The phylogenetic course of events is hard to disentangle; we
might suppose it to have been somewhat as follows :—
(1) Owing to the need of a firm support for pairing, it became
necessary, as above set forth, for the female to take the initiative
in the act of pairing, by assuming a special position.
(2) The male had thus no means of expressing his readiness
to pair [whereas in most monogamous birds it is the male, as
one would expect, who takes the initiative: cf the Warblers
(Howard, ’13), the Redshank (Huxley, ’12,), ete.].
(3) Meanwhile, quite independently, a process, or tendency —
eall it what you will—had shown itself, by which the characters
of one sex might be or tended to be transferred to the other, and
vice versa.
(4) This was seized upon by Selection (we cannot as yet speak
less metaphorically) and employed to supply the present want ;
the pairing attitude of the female was transferred to the male to
give him, too, a means of expressing his readiness to pair—to
enable him, should he wish it, as well as the hen, to take the first
step towards the performance of the act of copulation by the pair.
(5) As so often oceurs, the process did not stop precisely at
the desired spot (we still speak in metaphors, for brevity’s sake) ;
with the female pairing-attitude was transferred the female
pairing-instinct, and so came about the complete or nearly
complete facultative reversal of the pairing habits. ;
This naturally does not pretend to be more than a possible
scheme ; but it is worth while setting out such a scheme, merely
. aa how this ‘reversal of the sexes” could have come
about.
(ii.) Courtship.
T have started with the subject of coition, because the first
thing I want to make clear about the courtship-actions is their
total lack of connection with the act of pairing itself—a notable
fact, in which the Grebe differs radically, of course, from many
other birds, especially those in which the sexes differ in appear-
ance, é.g. the Bustard or the Peacock, but also some in which the
sexes look alike, e.g. the Redshank.
COURTSHIP OF THE GREAT CRESTED GREBE, 509
In relation to this, no doubt, is the fact that pairing only takes
place on the nest, and that the nest is hidden away among the
reeds, while the courtship actions are, I believe, aly ays gone
through out on the open water. This, i in itself, ‘would not be
conclusive evidence of total separation of the two sets of actions,
for the performance out in the open might be followed directly
by a veturn to the reeds and subsequent pairing. But there are
two further facts which make it conclusive. In the first place,
one of the reservoirs at Tring is completely bare of reeds, and con-
sequently of Grebes’ nests too. Itis, however, the richest in fish,
and numbers of Grebes fly over to it from the other reservoirs
every day, and at all hours of the day, to feed. Now, in spite of
the absence of reeds, and so of nests, and so of the possibility of
pairing, the birds interrupt their fishing, or sleeping, or preening,
to go through the ritual of courtship just as often on this
reservoir as on any of the others. ‘That is point number one.
Point number two goes still further.
I frequently kept individual pairs under observation for a
considerable length of time, and then, if I watched long enough,
always found that one set of courtship-activities would in point
of fact be followed by a pretty long interval of resting or fishing,
and that then this time spent in every-day affairs would be
again succeeded by another series of courtship-actions—a proof
that these actions are what we may call self-exrhausting and not
excitatory. The best record, because the longest, was on this
same reedless reservoir. J had one pair under observation for an
hour and forty minutes (section 10, record 11). During that time
they had six simple bouts of shaking, and also two prodigious
long bouts, followed each time by the diving for weed and then
the strange Penguin-dance. And between all these elaborate
displays of sexual emotion, no sign (or possibility) of pairing—
nothing but swimming, resting, preening, and feeding.
I was thus—much against my preconceived ideas—driven to
think of all the complicated postures and evolutions of courtship
in the Grebes as being merely an expression of emotion.
The particular form of expression used is no doubt determined
—predetermined—by the arrangement and innervation of certain
structures which the birds possess: but the impulse to use the
muscles and nerves is an emotional one—during courtship there
must be in the mind of the bird an excitement, a definite feeling
of emotion. Let us, to satisfy the physiologists, try to put it m
terms of nerve-currents. One member of a pair is continually
seeing its mate at its side. This, in its present physiological con-
dition, stimulates certain tracts of its brain, charging them up and
up until they are in a state of considerable tension (mental accom-
paniment:—state of diffused emotional excitement). Finally,
the tension reaches the critical point, and a discharge follows.
This discharge flows down hereditarily-determined paths, and
actuates the muscles concerned in courtship (mental accom-
paniment :—violent and special emotion, quickly dissipating
36%
510 MR. J. S. HUXLEY ON THE
itself with a sense of ‘“ something accomplished, something
done.”)
This merely indicates the possible material mechanism ; of the
actual, we know next to nothing. However, by comparing the
actions of the birds with our own in circumstances as similar as
ossible, we can deduce the bird’s emotions with much more pro-
bability of accuracy than we can possibly have about their nervous
processes: that is to say, we can interpret the facts psychologi-
cally better than we can physiologically. I shall therefore (with-
out begging any questions whatever) interpret processes of cause
and effect in terms of mind whenever it suits my purpose so to
do—which, as I just said, will be more often than not.
Let us take the parallel from human affairs. Far be it from
me to go into the matter with a heavy hand; let us merely
look at a few familiar facts in an unfamiliar biological light.
The “courtship-actions” of man are mostly predetermined by
heredity : any young couple that you like to take will be pretty
certain to ‘express their emotion” by holding each other’s hands,
by putting their arms round each other’s waists, or by kissing
each other; and of this last action kissing on the mouth is the
‘highest development.” Let us merely notice that these actions
are not perhaps exactly parallel with what we find in the Grebe—
that they are altogether more fluid, less fixed, and that they are
sometimes less self-exhausting and more excitatory in character:
on the whole, however, they are not very different. Moreover,
in thei case we know a great deal about the accompanying
emotions, either from our own experience or from what others
tell us. ‘To take only the most specialized form of human court-
ship-actions, the kiss; although we know that it may act as an
excitant (¢f. Dante’s famous lines on Paola and Fra ncesca) yet the
accompanying emotion is in itself quite special, different from all
others, and the emotional process is usually something an wnd fiir
sich, expressing itself in the action, and exhausting itself in the
process with a feeling of inevitability. In the memory, however
it, leaves its trace, and as it were desires to repeat itself, but
only when the emotional tension shall again have risen (think of
Plato’s epigram to Agathon: or the lovers in Richard Feyerel ;
or Romeo and Juliet). That will suffice to show what I mean by
a self-exhausting expression of emotion. Such a process would
be one that to the doer of it feels at the time almost inevitable
though he can only do it at certain moments. At other times.
determined by his general mental state (cf. section 10, record 1),
the action, however pleasant to recollection, is not “ spon-
taneously” possible, and if performed is forced or at least not
fully pleasurable. When normally executed, the action is accom-
panied by violent and pleasurable emotion, which usually dies
down, or changes, into a quite different feeling, one of satisfaction
meanwhile leaving its mark in the memory. Its recollection
then acts as a partial stimulus, so that next time it is a little
more easily performed.
COURTSHIP OF THE GREAT CRESTED GREBE. 511
This will, in the first place, show how difficult, and almost
inevitably futile, it is to try and deal with the emotional essence
of things by the methods of “ordinary biology”; I think, how-
ever, that it will serve to explain what I mean by a self-exhaust-
ing expression of emotion, and will give the point of view from
which to look at the facts of the Grebe’s courtship: let us now go
on to examine the facts themselves more systematically. I will
take the different forms of courtship-action one by one, describe
their usual occurrence and their relation to other actions, and
then mention the most important variations or exceptions that I
have seen.
The various attitudes already described are combined into
definite actions or ceremonies.
(a) The simplest form of courtship-action is the bout of shaking,
of which I have described a typical example. As already seen,
shaking may take place either before or after other courtship-
actions, but in perhaps the majority of cases it is not thus a link
in a chain of processes, but a single self-originating and self-
exhausting process. It varies acertain amount in intensity and in
length, and also in the amount of habit-preening that takes place.
Of this there may be none, or, towards the end of a bout, there
may sometimes be more preens than shakes. The bouts seem, to
the casual onlooker, to start themselves—in reality, I think, each
bird excites the other. One gently shakes its head under the
force of rising emotional tension; the other bird had not quite
got to that stage, but the sight of its mate shaking acts as a
stimulus, and it too pricks up its head a little and gives a shake.
This reacts on the first bird, and so the excitement is mutually
increased and the process fulfils itself—a very good example of
“‘ crowd-psychology,” and also a good example of an epigenetic
process *,
There is one well-marked variation of this form of courtship
which seems to denote a higher level of excitement ; it is especi-
ally common when a third bird has intruded into the domestic
harmony of the pair and has been driven off (section 5, iv.).
Here the beaks are pointed somewhat downwards, the neck
brought a little forward instead of vertical, the whole head
brought forward and curved over, and the ruff erected more than
usual (fig. 6). This attitude is almost always confined to the
beginning of a bout, the birds sooner or later relapsing into the
ordinary position.
The bout of shaking is not only the commonest form of court-
ship-action, but it also forms part of all the other more elaborate
forms. It always ends the series of actions, and often begins
them as well. It is as it were the foundation on which they are
built, and was probably (if I may express a mere opinion) the
earliest to appear in phylogeny.
The other ceremonies of courtship are all formed by the
* See also p. 544, where one bird won’t shake, and the other wants to.
512 MR. J. S. HUXLEY ON THE
combination, in various arrangements, of the shaking-bout, the
Dundreary, the Cat, the Ghost-dive, the weed-trick, and the
Penguin-dance; in addition, they may be slightly modified by
jealousy. ;
They can be divided into two groups: (1) those in which the
Cat-position plays a prominent part, and (2) those into which
weed-carrying enters. Let us consider them in this order.
The Cit-position forms a part of two quite distinct ceremonies,
which, simply for the sake of ready reference, I shall cali the Cere-
mony of Discovery and the Display. he first of these is gone
through, as far as I can make out, when the two birds of a pair
find and rejoin each other after being separated for some time.
‘The second always occurs in the middle of a bout of shaking; on
such occasions I presume that the shaking has not been ‘self-
exhausting,’ but that the emotional excitement that accompanies
it has reached a slightly higher level than usual, with the result
that it overflows into a new nervous channel, and so expresses
itself in this new way.
(b) The Ceremony of Discovery.—A typical case has already
been described (p. 497). I should interpret the facts thus :——The
two birds of a pair have become separated—perhaps they have
gone off fishing in different directions, or one has been on the
nest and the other has not stayed near by. ‘They wish to rejoin
each other. ‘l'o this end the bird that is searching puts itself into
a special attitude, which is probably adapted for uttering the
special ery only heard on such occasions, and cruises about, alter-
nating its signal-calls with moments of looking about it. On
hearing the call, several neighbouring birds will usually prick
up their necks and look about them; but I believe that it is
usually only the searehing bird’s true mate who takesany further
interest (this would doubtless depend on the emotional state of the
neighbouring birds), Once this discovery of the missing mate has
been made, a special ceremony takes place to celebrate the event.
This ceremony is a peculiar one, and is practically confined to these
occasions of discovery ; very possibly the memory of the ceremony
and its excitement adds to the eagerness felt by one bird of a pair
to rejoin the other. The ceremony itself usually consists in
this—the bird has been discovered dives, upon which the
searcher puts itself into what I have called the Cat-attitude, a
bizarre but beautiful position obviously recalling the elaborate
displays of many other birds. In this attitude the searcher waits,
almost always in a state of great excitement, as shown by its
turning itself hither and thither, from side to side. Jt is stimu-
lated to this excitement by the diving bird: first of all, as the
dive is very shallow, the diver’s approach is marked by a swift
vipple of the surface ; and then, when the diver at length appears,
it 18 in a shape as unlike that of everyday life as is the “ Cat-
position ” of the searcher—albeit the two are at opposite poles of
the Grebe’s capabilities, Sometimes the diver emerges when only
COURTSHIP OF THE GREAT CRESTED GREBE. 513
a few feet from the searcher. This is merely to reconnoitre his
position ; head and neck alone appear, the crest not erected, and
are swiftly withdrawn again, The final appearance takes place
almost always beyond the searcher, and the bird emerges with its
back to the other, facing it only as, revolving on its axis, it settles
down. The performance always ends with a bout of shaking.
Although in my “typical” cases the searcher has always been
the female, yet the male may also search for his mate in the same
way. I have watched an obvious cock in the regular Dundreary
(search) attitude for a long time; only on this occasion no mate
responded to the call. Further watching is necessary to see
whether it is merely an accident that my searchers have usually
been hens, or whether my observations represent reality. (It may
possibly be connected with the fact that the hens seem to spend
more time on the nest than the cocks; but this is mere con-
jecture. )
(c) The Display Ceremony is quite different. The birds are
already together, and the display is simply a form of excitement
similar to the bouts of shaking. In typical cases the pair will be
indulging in a bout of shaking; suddenly one of them flies off a
few yards and puts itself into the full Cat-position, showing its
circular ruff and white-striped wings to its mate. There is no
diving, however, and, after some seconds’ display, the birds swim
together and there is another bout of shaking; after this they
simply swim off, or separate, or feed. Hither eock or hen may go
‘into the Cat- attitude. In one ease the first bout of shaking ‘had
been preceded by a “ flirtation” (p. 521) on the part of the cock.
To show how one ceremony may blend into another, I adduce
the following instance :—There was a regular Discovery Ceremony,
the hen calling to the cock, but with this difference, that they
swam together, shook, and the hen flapped off and went into the
Cat-position, oral that only then did the cock remember, so to
speak, to do the ghost-dive (p. 498). Another mixed “cer emony,”
this time more closely related to pure display, is related on p. 547.
(2) Finally comes the ceremony of the Penguin-dance. I have
little to add to the description already given (p. AQ9): Wwice,
curiously enough, a single pair was seen to perform the dance
‘twice In a morning. This might imply that some special physio-
logical state, pr obal ly of high excitement, was necessary for the
act, ror I only saw it on two other occasions, and Selous only saw
it once.
The performance can only be gone through when both
are equally excited; for instance, once (p. 547) after a bout of
shaking the cock dived and fetched weed from the bottom. The
hen, however, was not stimulated to do so too, and when he came
up he found no answering stimulus, and so dropped the weed as
he swam towards his mate.
There is no reason for supposing even this elaborate ceremony
to have any direct relation whatever with coition. It is a form
5IL MR. J. §S. HUXLEY ON THE
of excitement and enjoyment, seemingly as thrilling to the birds
as it is to the watcher, but, like all the other courtship-actions,
self-exhausting. 3 :
Very interesting “‘incomplete stages In development of this
ceremony were seen in one pair of birds, ranging from simple
diving to the complete ceremony (see below).
(e) Other Courtship Ceremonies.
(1) Back-to-back Ceremony.
There are considerable individual variations in the courtship-
activities of the Grebe, and I have seen occurrences which may
well be interpreted as rudiments of new ceremonies. In one pair,
for instance (section 10, record 12), the birds almost always went
into a formal back to back, or rather tail to tail, attitude after
each bout of shaking.
(2) Diving Ceremony.
Other actions seem to stand in some relation to the more highly-
developed ceremonies. For instance (pp. 545, 552), simple diving,
either by one or both birds, without any fetching of weed from
the bottom, is introduced as part of the courtship between the
two bouts of shaking.
(3) Weed-trick Ceremony.
In still other cases, the weed-trick is gone through and is
followed immediately by a bout of shaking. In both pairs in
which this was seen, one bird alone brought weed, although in
one pair (not in the other) both had previously dived. I am
inclined to believe (but more observations are needed) that two
(distinct ceremonies are here involved: first, the fetching and
offering of weed by one bird, usually the cock, to its mate; and,
secondly, a typically “mutual” ceremony involving simultaneous
diving of both birds of a pair: and on to this latter the “ penguin
dance” has been grafted.
The offering of weed is strongly reminiscent of occurrences in
the sex-differentiated, non-mutual courtships of other birds, such
as the Warblers, where the cock often carries Jeaves or twigs
in his mouth during sexual ecstasy (Howard). It seems to me
probable that, since diving is necessary for weed-fetching, the one
has come to be associated with the other, and the two ceremonies
have come to be mixed up: in the extreme case on one side there
is no mutual ceremony—only an offering of weed by one bird to the
other; at the other extreme we have the complete penguin dance
as described on p. 499, where both cock and hen bring up weed ;
and as intermediates we have mutual diving (p. 552) and mutual
diving where the cock alone brings up weed (p. 552, end).
Taken all in all, the courtship is chiefly mutual and self-
exhausting: the excitatory, secondary-sexual forms of courtship
such as weed-offering or pure display serve not as excitants to
COURTSHIP OF THE GREAT CRESTED GREBE. 515
coition, as in most birds, but as excitants to some further act of
courtship ; and this is always a mutual and also a self-exhausting
one. ‘The excitants to coition are of a very special nature, and
are symbols, rather than mere general excitants.
Habit-Preening. (See p. 497.)
This is very frequent, occurring in about half the bouts of
shaking seen. The more excitement, the less preening, seems to
be the rule ; long bouts may sometimes degenerate into practi-
cally undiluted preening, the head simply being brought more or
less up, but not shaken, between the “ preens.” It is always the
hind end of the wings, I believe, which is raised and let fall by
the beak.
In some way there must be a strong association between
preening and head-shaking in the Grebe, for solitary birds who
were really preening themselves I have several times seen raise
their heads, slightly bristle their crests, and give a rudimentary
shake. Why or how the association has taken place is more
difficult to say. I certainly believe that the action I call habit-
preening has been derived from true preening, and has been
ceremonialized in the process of becoming part of a courtship-
action. For the present we must leave it at that,
‘© Habit-Shaking.”
That for some reason there is a very real association between
shaking and preening is shown by the following facts. When
actively engaged in real preening of themselves, the birds are
often seen to lift their heads, give a rudimentary shake or two
(without erecting ears or ruff) and then go back to business.
This is generally seen when the bird is engaged in preening its
hinder parts. We have observed it in autumn as well as in
spring, and so it presumably takes place the year round ; there is
thus obviously a real association between the preening and the
shaking, and the shaking is not a mere release of simpler sexual
energy.
This is exactly the converse of what I have called ‘habit-
preening,” and may therefore appropriately be styled “ habit-
shaking.”
There is thus a single association with a two-fold result. How
it can possibly have arisen, or what purpose it can serve, remains
to me at present an absolute mystery. I leave it as a puzzle to
future bird-watchers and comparative psychologists.
Fighting between Cocks.
I saw very little of this beyond mere hostile expression (p. 521).
Once, however, I saw two birds actually grappling: one was
struggling half-submerged, while the other was more or less on
top of it, and had hold of the feathers of the back of its opponent’s
head. After some considerable splashing and struggling, they
separated and swam apart.
516 MR. J. S. HUXLEY ON THE
In birds which pair up early and remain “ married” for the
season, hike the Grebe, one would, of course, not expect to find any
of the regular combats seen in other species. It would be inter-
esting to see whether there is more fighting in February, during
the actual process of pairing-up.
The question must now be put—‘ What for?” What is the
good of all these divings and posturings, these actions of courtship,
these ‘‘expressions of emotion”? ‘lo what end are colours and
structures developed solely to be used in them, and what return
is got for the time and energy spent in carrying them out ?
They are common to both sexes, and so have nothing to do with
any form of true sexual selection; they are self-exhausting pro-
cesses, not leading up to or connected with coition, and so cannot
be sexual excitants in the ordinary sense of the term.
It must be, however, that they fulfil some function; and I
believe I know what this function is. I believe that the court-
ship ceremonies serve to keep the two birds of a pair together,
and to keep them constant to each other.
The Great Crested Grebe is a species in which the two sexes
play nearly equal parts in allactivities concerned with the family.
The cock shares equally in nest-building, -nearly eyually in in-
cubation and early care of the young (it is only later that the
young pass into the care of a single parent, probably the female,
see Pycraft,’11). Thus, from the point of view of the species,
it is obviously of importance that there should be a form of
“marriage ””—a constancy, at least for the season—between the
members of a pair. ‘The same result—marriage—is observable in
such a species as man; but in man the main cause is a division of
labour between male and female, whereas in the Grebe the sexes
have been made as similar as possible. It would seem that the
Grebes’ family affairs had simply required more labour to be spent
on them, and that Evolution had happened to go along the simple
path of increasing the quantity of labour, by bringing the male
in to do female’s work, instead of improving the quality by
adopting the principle of specialization.
Birds have obviously got to a pitch where their psychological
states play an important part in their lives. Thus, if a method is
to be devised for keeping two birds together, provision will have
to be made for an interplay of consciousness or emotion between
them. It would be biologically enough if they could both quite
blindly, and separately, attend to the common object—nest,
eggs, or young; but with brains like theirs there is bound to be
_a considerable amount of mental action and reaction between
them. All birds express their feelings partly by voice, and very
largely by motions of neck, wings, and tail; and not only this,
but the expression can be, and is, employed as a form of language.
This being so, we have here a basis on which can be reared various
emotional methods of keeping birds of a pair together. As
always, selection of accidental variations has led to very diverse
results ; so that we see this ‘‘ emotional companionship ” playing a
COURTSHIP OF THE GREAT CRESTED GREBE. 517
art in many apparently very different actions of birds. Herring-
Gulls sit or stand close beside each other for hours together,
occasionally rousing themselves to a joint ceremony of shaking
their necks. As the Snipe drums overhead, there is often a call
fromthe marsh below. Many birds when paired are always calling
to each other, and probably singing birds sing partly to their
mate; Dabchicks have a special spring note, usually given as a
duet. As a very simple case, I have seen a pair of Blue Tits
very recently paired up who, although feeding, were perpetually
ealling to each other and at frequent intervals coming close up
side by side; it was perfectly obvious that they simply took
pleasure in each other’s presence, like the engaged couple that
they were.
We have thus the following train of reasoning. Many birds
must be kept in pairs during the breeding-season. ‘This may be
partly effected by the instincts of the separate birds—the mstinct
to build a nest, to sit on eggs, to feed young; and partly by
sustincts which only can find play when the two birds are
together. ‘These latter are often very emotional, and the court-
ship habits of the Grebe afford a very specialized example of this
emotional bond between members of a pair.
1f my contention is correct, it is clear that many actions and
structures solely used in courtship are of use to the species, and
not only to one sex of the species; these therefore must be
maintained by Natural as opposed to Sexual Selection,
(iii.) Nest- building.
I rely almost entirely on the observations of Selous (01, and
see my Summary, section 8).
Materials —Selous’ nest was made mainly of dark ribbony
weed, fetched from the bottom. Some surface-weed was also
used. Besides this, such objects as water-lily stalks and large
water-logged sticks are occasionally employed, the latter seeming
to help anchor the nest. To steady the nest still further, the
weeds are often woven among the stems of growing water-plants.
As with other birds, the materials vary with the situation. In
Avundo-heds the nest is very largely made of bits of reed-stem,
though always with some weed; while elsewhere no reed is used
at all.
Time, etc—The main bulk of the nest appears to be built very
quickly—in a few hours, in fact. This main portion is always
built in the early morning (as with many other birds), and while
on one day there may be no nest to be seen, by the next it may
be nearly completed. In this very active building both cock and
hen often take part simultaneously; they work very hard,
averaging between them more than two cargoes of weed every
minute, and gomg on for half an hour or an hour at a time
without stopping.
After one nest has been built, another may be started (and
518 MR. J. 8. HUXLEY ON THE
nearly finished) next day at no great distance. In the case
noted by Selous, during the building of each nest the cock
built himself a platform or rudimentary nest close to the bank
and not far from the nest that was in progress. In the con-
struction of this the hen did not share.
FUNCTIONS OF NEST.
There seem to be at least three kinds of nests—the true
nests, the pairing platforms, and the cock’s platforms.
(1) The true nests are bulky structures, rising well out of the
water. A single pair of birds may build more than one. In one
of these the eggs are finally laid. It is not known whether a pair
always build more than one (probably not), nor whether, if more
than one is built, it is always the last that is chosen to receive
the eggs.
(2) The pairing platform (one seen by Selous and one by me) is
a sodden, messy-looking structure, apparently much trodden
down, and practically flush with the surface of the water. It
seems to be used only as affording the necessary support for the
action of pairing, and is apparently built some time before the
true nest, e. g. Selous (01, p. 339 e¢ segg) watched a pair for
three weeks and saw no nest except a single platform, which
was used exclusively for pairing (and pairing actions). It is
distinctly improbable (from the appearance of all true nests that
I have examined) that such a platform would be built up and
turned into a true nest, though no evidence on this point is
forthcoming.
What is certain, however, is that pairing need not always take
place on such a platform, since Selous saw it occur ona true nest,
and one which had probably one egg already in it. It appears to
me likely that such platforms are built early in the season, when
only copulation (or the preliminaries thereto), and not incubation,
isin progress. Then, later, true nests are necessary to lay the
eggs in, and once these are built copulation can take place in
them just as well as on the platforms—. e. the platforms are only
temporary expedients, rendered necessary by the birds’ pairing
habits, and would thus phylogenetically appear to be degenerate
nests.
(3) The platforms of the cock. These, too, are imperfect
structures, and are probably also degenerate nests. In the case
observed by Selous they were built close to the bank by the cock
alone, during the construction of a true nest close by. When
built, they were used (by the cock alone) as resting-places where
he sat or, more frequently, stood.
Their true purpose is hard to see. It cannot be that the cock
needs a resting-place more than the hen (unless—merely to state
a case !—we suppose that in the month of May both sexes need
COURTSHIP OF THE GREAT CRESTED GREBE. 519
to rest on some firm support more than at other times, and that
the hen incubates more than the cock, so that he has to build
himself an extra resting-place). Possibly such platforms have
something to do with receiving the young after hatching, and
are built precociously. But this is mere guesswork—further
watching must reveal their secret.. Mr. Selous, in a letter to me,
says he thinks now that the platform is due to an aberration of
the nest-building instinct inthe male. From Selous’ own records,
however, it is clear that the male certainly uses it to rest on.
Differences in building of the different kinds of nest.
The bulk of the true nest is built by both birds together in a
very short time. After that, it seems to be casually added to
during the time of incubation. his is useful, for the nest would
otherwise get gradually pressed down into the water. It also
tends to lose its slightly cupped form, and to remedy this the
sitting bird may often be seen to pick bits of weed from the inside
or the outside of the nest and lay them on the rim.
The cargoes of weed brought during incubation are often very
small compared with those brought during the first building of
the nest, and the whole action seems often to have rather the
stamp of a habit about it—“ I am going to the nest, so perhaps
I had better bring a weed or two along.”
During the main building, one bird sometimes sits on the nest
fer a short time ; during this period one bird might also lie along
the nest in the passive pairing position, as an invitation to the
other to pair. But apparently pairing and pairing-actions are
gone through more often when the bulk of the nest is finished,
or when a platform alone exists—nest-construction thus ap-
pearing to use up most of the emotional energy of the birds.
The cock’s platform.—It would be interesting to know if this is
always made during the construction of the main nest, or if
sometimes the cock set about it in cold blood, before or after the
true nest was finished. It is quite possible that the cock and
the hen stimulate each other to active nest-building as they do to
courtship activities (p. 511), and that while they are in this
excited state, and only then, some of the nest-building energy is,
in the cock, diverted to his platform.
The pairing platform.—This (only one recorded case) was
already built when Selous first noticed it. Only small pieces of
weed were added to it, and usually “in a very perfunctory
manner” (Selous, 01, p. 342). After one bird had unsuccess-
fully invited the other to pair, it often ‘‘manipulated the weeds
a little with its bill” before coming off the platform.
We thus badly want to know about the building of the pairing
platform; and also whether, during its construction, the cock
builds a platform for himself, as he does during the building of
the true nest.
Or
i)
oO
MR. J. S. HUXLEY ON THE
Share of the sexes in nest-building,
The only data are based on the actions of a single pair—that
observed by Selous in 1900; we must not, therefore, generalize
too far. However, certain things emerge clearly. Both sexes
work vigorously at the nest. During nest-building, the cock
builds a platform by and for himself not far from the nest, In
this particular pair the cock seemed slightly more active in nest-
building, though the hen was more skilful. This may be, and
probably is, merely an individual trait. As far as the building
of the true nest goes, both sexes seem to be at least as similar as
they are, for example, in appearance and in courtship habits.
‘The mysterious platforms of the cock remain as one of the few
truly secondary sexual characters of the species.
Psychology.
It seems probable that the same sort of psychological mechanism
holds here as in courtship. Fetching weeds from the bottom and
piling them in a heap is an instinctive act affording pleasure or
relief. Although it is much easier to Imagine a bird deriving
pleasure from solitary nest-building than from solitary courtship
action, yet here, too, it seems as if the cock and hen are mutually
stimulated to activity. This is, at all events, an eminently useful
trait, for it ensures that the nest shall be quickly built, that
the pair shall keep together, that they shall build one nest at
a time (instead of two, perhaps widely separated ones), and so
forth.
That it is pleasurable may be further inferred from the fact
that when the fever is upon them the birds may build two large
nests in two successive days (Selous, loc. cit). This may seem
wasteful, but here, as in so much else, Nature indulges in a
considerable reserve—better too much than too little. This is
the case with reserves of food, our own appetites, the number of
times which most animals pair, the number of unpaired males
in polygamous species and in Bees, etc., ete. Here, perhaps, the
reserve is indirect rather than direct; it may not even be any
advantage to have built two nests instead of one, but it may well
be an advantage to have such a strong nest-building instinct that
two or more nests happen to result, instead of one.
Finally, again, as in the courtship, association plays its part.
What has once been done with pleasure, is done again easily as a
habit. So when a bird is near the finished nest. and especially
when the birds are there together, weeds are often added, but
almost always in a more or less perfunctory way.
Why, after one bird has unsuccessfully invited its mate to pair
on the platform, it should often dive and bring weed to the plat-
form, I donot fully see. We can only say that diving and weed
COURTSHIP OF THE GREAT CRESTED GREBE. 521
are connected by separate and perhaps roundabout mental paths
with nest-building, courtship, and pairing; and that what we
know is only an outline of the birds’ behaviour.
(iv.) Relations of different pairs to each other.
(Fuller details are given in Part J5))
The Great Crested Grebe, as we have already seen, pairs up very
early in the season. What is the relation of the pairs of birds
to one another ¢
Asa general rule, the two birds of a pair seem to take very
little notice of the rest of their species. Oceasionally, however,
there is some contact. There may be jealousy on the part of one
bird, and this jealousy may, in any particular case, be merely
precautionary, or actually justified by flirtatious behaviour on the
part. of the other bird of the pair; or there may be hostility
between members of one pair and members of another.
Simple hostility is the rarer of the two: the only reason I can
discover for it is the trespassing of one or both birds of a strange
pair upon the * territory ” of another (p. 558). In its symptoms
there is nothing very remarkable: the birds go into the threaten-
ing Dundreary-attitude, often “barking” angrily at each other ;
finally, one may fly or dive at the other and drive it away, but
often the very mild form of hostility involved in staying quite
still, assuming the threatening attitude, and barking at the
enemy (who is also doing the same) is all that happens.
Tt is much more interesting, however, to find in these birds
what we had best call flirtation, aS an accompaniment to their
monogamy. The whole thing is very human : when one member
of the pair is rather excited and the other is either lethargic or
far away, there 1s no channel for the relief of its excitement. If
a bird of opposite sex is in the neighbourhood, however, this
would provide the desired relief, and the result is that the
“temptation ” is often too strong, and a bout of shaking ensues
between two birds who are not mated. J have never actually
seen such flirtations go beyond a bout of shaking; there is no
yeagon that they should not, except that, as far as I could see,
the birds did not seem to be used to each other, so to speak,
Zand so their excitement often cooled down very quickiy.
Tf the rightful mate sees what is going on, it is always
roused to action, however lethargic it may have been before.
Tt drives the odd bird away (often by a subaqueous attack with
the beak) and then almost always has a strong pout of shaking
with its mate. Thus all the anger of jealousy 1s directed against
the usurper, not against the mate—which again 1s distinctly
human! ‘The “erring spouse "is always equally ready to shake
with his mate as with the tertium quid—and often more so.
Here, again, the sexes are qualitatively alike—either will take
522 MB. J. S. HUXLEY ON THE
the initiative in flirtation, although from my records the cock
seems to do so rather oftener than the hen.
Where there is a simple pleasurable ceremony, for whose
performance two birds are necessary, it would seem quite natural
that flirtation would occur. If the ceremony is an advantageous
one, flirtation represents an overshooting of the mark by Natural
Selection—a slight disharmony. ‘‘ Adultery” I would think very
improbable in this species, since the act of pairing is connected
with a nest, built jointly by the pair, in a definite spot of their
own territory.
(v.) Other Activities.
Incubation.
It appears that both sexes sit, but that probably the hen sits for
a much longer time. Mr. A.'T. A. Ritchie informs me that when
there is a punt near the nest and the hen does not want to return
to the eggs, the cock will often drive her to her duties.
Care of the Young.
After the young birds are hatched, both parents attend on
them for some time; but later in the season, when the young are
half-grown, the observations of Mr. W. P. Pycraft (Pyeraft, °11)
make it certain that only a single bird, probably the hen, looks
after the brood.
Thus here there is no complete and qualitative division between
the sexes, except in this last particular.
There is, however, rather more of a quantitative division than
usual.
6. Discussion.
There are various considerable difficulties concerned with the
courtship-structures and actions of the Great Crested Grebe.
In the first place, it is clear, from what has been said, that in this
bird there is no sexual selection in the ordinary sense of the
word ; the crest and the courtship-actions are almost identically
developed in cock and hen alike.
On the other hand, the crest is only fully developed in the
breeding-season, thus resembling true secondary sexual characters;
and, as I have pointed out (Huxley, ’12,) it is used only in
courtship, so that if not “secondary,” it is at least “sexual.”
Further, the crest is smaller (though but slightly) in the
female than in the male, a fact which it is, at first sight, simplest
to explain by assuming that the crest was acquired by the cock
as a secondary sexual character, and has now been almost com-
pletely transferred to the hen (cf. similar transference, complete
or incomplete, in Lycenid and other butterflies (Weismann),
Reindeer, mammz of mammals, colours of many birds). We
will revert to this point.
COURTSHIP OF THE GREAT CRESTED GREBE, 523
The courtship actions, however, can scarcely be explained by
transference. ‘The Penguin-dance, for instance, can never have
been anything but a joint ceremony, equally shared by both sexes.
Furthermore, even in the Dabchick, although it (and it alone in
the subfamily) lacks all courtship-structures on the head, there is
a joint courtship-action—the two birds come face to face, stretch
up their necks, and emit the well-known cry. This being so, it
is fairly clear that the ancestral courtship-actions of the Grebes
were not in the nature of a display by one sex, but were joint
actions of the pair. ‘There is nothing especially remarkable in
this. The display-courtships are, on the whole, more striking,
and so have been more frequently described ; but (to draw on my
own limited experience) Razorbills.and Herring-Gulls have very
well-marked joint courtship-actions, although the actions are
associated with no special structures whatever, and Selous has
described other such actions in Swans, Divers, Guillemots,
Fulmars, and other species.
I should put forward the theory that the courtship-habits of
birds are based upon at least two totally different foundations :
in the first place the actions gone through by males alone, appa-
rently as the direct result of sexual eagerness (solitary actions),
and, in the second place. the actions gone through by male and
female together, and perhaps often (though by no means always)
connected or associated with nest-building (combined actions).
Primitively in neither case would there be any special structure
or colour associated with the action. For solitary actions this is
well seen in the dowdy Warblers, so fully described by Eliot
Howard; here the cocks resemble the hens, but go through
elaborate droopings of wings and fannings of tail, with bristlings
-of feathers on throat and crown. Later, Sexual Selection has
stepped in, and naturally enough has taken what was already
given, and added to it. The same instinctively-displayed parts—
wings and tail, throat and crown—are the parts which are
especially singled out for the development, first of special colours
(Finches, Woodpeckers), then of special colours and structures
combined (Turkey, Argus Pheasant, Blackcock, etc.). In com-
bined actions a similar process has been gone through. In the
Herring-Gull and Razorbill we have the instinctive actions pure
and simple—a direct outcome of nervous excitement. Then,
again, something has stepped in and used what was thus provided,
and we get combined actions displaying colour (coloured mouths
of Fulmar Petrels, Selous), and finally colour and structure, as in
the Grebe. The members of the Heron tribe in general, and the
Egrets in particular, have also ornamental structures common
to both sexes; it would be very interesting to know the course
of courtship in these birds. Pycraft (13) figures a mutual dis-
play executed by the Kagu.
The question now arises, How have such colours and structures
arisen? By Sexual Selection followed by transference, or by
Proc. Zoou. Soc.—1914, No. XXX VII. 37
524 MR. J. S. HUXLEY ON THE >
some other process? Such other process can easily be imagined,
and I feel confident that it has played a considerable part. We
may call it Double or, better, Mutual Sexual Selection (Mutual
Selection for short). Where combined courtship-actions exist,
and a variation in the direction of bright colour or strange
structure occurred, it would make the actions more exciting and
enjoyable, and those birds which showed the new variation best
would pair up first and peg out their “territories” for nesting
before the others could get mates. The level would tend to be
raised generation by generation. Mutual Selection is in a way
a blend between Sexual and Natural Selection. The structures
and actions arising under it have their immediate origin in the
preferences of individual birds, not in anything outside the
species, and in their immediate function they are entirely confined
to the courtship. On the other hand, the mutual courtship
itself, the activities of both birds taken together, may be of use
to the species as a whole, in keeping the sexes together when
necessary. ‘Then the indirect function of all the shaking-bouts
and displays of the Grebe is a function of use to the species, and
besides the direct origin there is added an indirect origin under
the pressure of Natural Selection.
Mutual Selection has a certain similarity with assortative
mating, but is by no means the same thing. Like true Sexual
Selection, it encourages an ever higher level in the development
of a character, once variation has given it a basis to start from.
In the Grebe the line of variation encouraged by Mutual Selection
has been the tendency to produce ruffs and tufts of feathers on
the head, and to go through actions involving, besides the use of
these structures, diving and sporting with water-weed.
The question in the Grebe is complicated, as noted above, by
the slightly less developed crest of the hen ; this, however, might
easily be accounted for by differences in the metabolism of cock
and hen. The Discovery and (especially) the Display Ceremonies
are also rather stumbling-blocks in the way of an explanation by
Mutual Selection ; they seem so very like the Displays of solitary
courtship. However, even here the second bird plays a part,
which in the Discovery Ceremony is at least as important as that
of the displaying birds.
What is quite clear, however, is that, even supposing (what
to me personally appears very doubtful) that ordinary Sexual
Selection has “produced” the structures and the cat-position
(we must know more about the habits of other species of the
genus to decide this), yet it has gone hand-in-hand with a
process of Mutual Sexual Selection as regards the majority of the
actions. These actions (like the display of the Peacock, but
unlike that of the Warblers) are much too elaborate and much
too specialized to be considered as the immediate outcome of any
form of physiological excitement. They obviously have a long
and complicated evolution behind them, and, as they can only
525
On
COURTSHIP OF THE GREAT CRESTED GREBE,
be performed by the two birds together, there is nothing to
account for them as they now stand but some such process as I
have just sketched under the name of Mutual Selection
Then there comes the question of the facultative reversal of the
act of pairing (or, possibly, only of preliminary pairing-attitudes).
The other cases noted by Selous (Pigeon and Moorhen) differ in
that the male crouched to the female directly after the act
of pairing, who at once proceeded to play the male’s part. In
the Grebe there was always a long interval before the ‘“ reversal
of instinct ” took place.
In all, however, it is very difficult to see how to account for it,
except on the assumption that there has been a reciprocal
“transference” of pairing-instincts. This transference may be
apparent or real. It is apparent if we believe that the units for
such sexual characters are equally present in the germ-plasm of
both sexes, and that the chavacters themselves do not appear in
the other sex (or only appear as rudiments) asa result of the
great primary sex-difference.
If the transference is real, then one must assume that the
zygotic constitution of the two sexes is different in regard to
secondary as well as primary differences, but that there is a
constant tendency—depending on some as yet unknown process—
to transfer such characters to the opposite sex. (Hybridization
experiments, where the female of a species can transmit to her
male hybrid offspring the secondary sexual characters of her own
species, indicate that the first method isthe true one.) How else
than in one of these two ways can we explain transference in
both directions? This is seen, for example, in man, where a male
organ, the moustache, appears rudimentarily in the female, and
female organs, the mamme, appear rudimentarily in the male:
in abnormal cases, besides, the transference may be complete, the
organs being completely developed in the wrong sex. Such
moustached women and men with breasts again support the idea
that the transference is not a real transference, but consists in
the removal of an inhibition only.
(L would not trouble to mention the theory that these appear-
ances of characters of one sex in the other are due to descent
from a hermaphrodite ancestor, were it not actually the case
that Metchnikoff has advanced it. It is enough to point out that
if this were so, the primitive mammal must have been a herma-
phro lite.)
To us it makes little odds whether there is inhibition alone or
transference followed by inhibition. In both cases the character
will be in antagonism with the inhibitor: supposing that there
is no longer any need to inhibit a character of one sex in the
other, then on Darwinian and Weismannian principles the in-
hibiting “force” will atrophy, and the character, remaining
as strong as ever, will appear equally in both sexes.
Apply this to the present case. Birds are for the most part
Bi
526 MR, J. S. HUXLEY ON THE
so constructed that impregnation would take place equally well
whether the sexes are in normal or reversed position : that is to
say, there is no necessity for keeping to the customary position—
and accordingly “reciprocal transference” of the pairing atti-
tudes (whether the transference be apparent or real) may, and
quite probably will, take place. If so, then in one of our Grebes
the instincts and reflexes for the pairing-actions proper to its
sex co-exist side by side with those for the pairing-actions proper
to the other sex. It is also obvious, first, that both cannot be
gratified simultaneously ; and, secondly, that these two very
different sets of actions must be associated with two very different
sets of emotional states. The bird may ‘“‘feel female” or it may
“feel male,” and according to its feelings, so will it tend to act.
But, as we saw before, in discussing the pairing-attitudes, it
appears that, owing to the difficulty of coition in the Grebe, the
“female” (passive) pairing-attitude has become a mere symbol
of readiness to pair. Thus Natural Selection has come in to
assist the slow process of transference (at any rate, so far as
pairing-attitudes are concerned), and since whatever involves
them will probably involve coition itself as well, we have an
additional reason for believing that actual reversal of pairing
does take place, as Selous supposes, in the Grebe.
At any rate, there can be no doubt about the reversal in the
Pigeon and Moorhen. The sudden reversal that here takes place
is rather different, but may be explained somewhat as follows :—
Here, too, both active and passive instincts are now represented
in either sex. A bird is ina state of sexual excitement ; this
excitement releases itself in the performance of, say, the male part
in the act of pairing. The excitement is not always completely
exhausted by the act, and, if so, the act is repeated (just as the
shaking-bouts of the Grebes are continued for a longer or shorter
time, according to the degree of what we may call courtship-
excitement). But supposing that general sexual excitement
arouses both the male and female emotional states, then the per-
formance of the act once in the male attitude will only exhaust
the feeling of ‘male excitement,” leaving the ‘female feeling ”
still a-tingle. The result will be, first, an inducement to repeat
the act and, secondly, an inducement to repeat it with attitudes
reversed.
Thus such immediate reversal is more or less an accident of
heredity, while the Grebe’s reversal is an accident aided by the
usefulness of the transferred actions, which thus bring the acci-
dent within the sway of Natural Selection,
This treatment of the question is of necessity sadly speculative,
but it is our duty at least to try to construct a coherent mechanism
of theory to explain the isolated facts of observation.
Finally, a word as to terminology. J have already pointed out
(Huxley, ’12,) that the phrase secondary sexual cannot be applied
to the Grebes’ ruff and ears or to their courtship-actions, because
COURTSHIP OF 'THE GREAT CRESPED GREBE, 527
this term always implies a difference between the two sexes, and
yet the crest of the Grebe has a sort of secondary sexual look
about it—unreflectingly, one would at once write it down as
such. This is due to our incomplete classification. We begin
by separating out sexual characters from all others—these being
characters that are different in the two sexes. We divide them
into primary, accessory, and secondary. The mammee of mammals
(with the exception of man) have nothing to do with courtship
or mating, yet they are usually included under the same heading
as the tail of the Peacock, while the Grebes’ courtship-structures
would be left out in the cold.
Besides the mere criterion of difference in the two sexes, we
must have some other criterion—a criterion of use.
It is naturally impossible to draw up any completed classifi-
cation that will satisfy every case. To do so would be beyond
the powers even of a Herbert Spencer—and not of much use when
done.
It is enough to point out, first, that our group of Secondary
Sexual Characters is a bit of an omniwm gatherum. Some of
them, as the mimicry of the female Papilio, or the brown colour
of the female Pheasant, are protective, of use to the individual
and to her offspring. Others, such as the mammee of mammals,
are of use only to the offspring; others, like the sexual differ-
ences in the beak of the Huia, where male and female hunt in
couples, one splitting open the wood, the other picking out the
hidden grubs, have arisen by a division of labour, and are of use
to the couple as a couple. One might go on, but it would be
unprofitable.
In the second place, we must recognize as a fact that the
existence of individuals of separate sexes with wills of their own
has led to the development of what we call courtship—simply a
process in which a series of actions is carried out as the outcome
of an emotional state based on sexual excitement. All courtship
is based on sexual excitement, and characters connected with
courtship merit a separate name of their own. This name lies
ready to hand in Poulton’s term epigamic; we must, however,
remember that the literal meaning of the term must not be
pressed, for in many cases the courtship ceremonies do not lead,
directly or indirectly, to the act of pairing. Let us rather turn
it the other way about, and, defining an epigamic character as one
that is used in courtship, go on to define courtship as a series of
actions based immediately or remotely upon sexual excitement,
and, to make ourselves clear, we must add that sexual excitement
is not merely sexual desire, but that whole emotional state into
which a member of one sex may be thrown by a member of
the other. The necessity for the distinction is obvious, if
we think of the conditions in Man. Sexual excitement, of
course, includes mere sexual desire, and also includes the fighting
of males among each other as a result of sexual desire.
528 MR. J. S. HUXLEY ON THE
~ Tf we want a tabular statement, we can draw up something
like the following * :—.
(A) Characters different in the two sexes.
(Sexual characters).
(1) Primary. Of the gametes and gonads.
(2) Accessory. Concerned with the union of the gametes.
[Copulatory organs, pairing attitudes,
sexual desire. |
(3) Secondary. All others.
(a) Developed through Natural Selection.
(Huia beak; mamme; marsupium ;
incubation by @ alone in birds, &e. }
(b) Capable of being developed through Sexual > Epigamic.
Selection. | (Courtship characters, i.e. all
[Horns of deer; tail of Peacock, &e. | characters concerned with
(B) Characters similar in the two sexes. r the selations of the sexes,
(1) Capable of being developed through Mutual | excepting those connected
Selection. immediately with coition.)
[Grebes’ conrtship and crest ; Herring-
Gull’s courtship, &c. }
(2) All other characters.
It might perhaps be better, as has been suggested to me, to
restrict the term Secondary Sexual to 3b, and employ Sex-limited
where I have employed secondary sexual. For one thing, how-
ever, this would conflict with Darwinian use; also, I am at
present more concerned to show the necessity for new thinking
than for new terminology, which will be more suitable in a more
general and definitive paper.
IT will conclude by hoping that anyone who has the oppor-
tunity will observe the habits of the Crested Grebe during the
time of pairing-up in early spring; the full courtship of the
Dabchick would also be of very great interest. In the near
future, I hope to publish a more general paper upon Mutual
Selection, so that any notes sent to me on this subject will be
gratefully received and acknowledged.
For some further discussion, I refer the reader to the Postscript
(p. 559).
PART II.
7. Locatiry, Mrtuops, Erc.
Through the kindness of the Hon. Walter Rothschild, I was
given a permit to watch, and a punt to watch from, at the Tring
Reservoirs. These consist of four large sheets of water in the
eastern corner of Hertfordshire. Two have dense beds of reeds
* Simple sexual desire, if we adopt this scheme, we may call an accessory sexual
emotion. Sexual excitement includes this, and all the epigamic sexual emotions as
well. Pairing attitudes are accessory sexual attitudes. Let it, by the way, never
be forgotten that emotions and attitudes are just as much characters as are colours
or structures.
COURTSHIP OF THE GREAT CRESTED GREBE. 529
(chiefly Arundo phragmites) along one shore, affording cover for
the nests of the Grebes, while the banks of the other two are
bare. One of these latter is, however, very rich in fish, and a
number of Grebes come over every day to feed on it. There,
of course, they are nowhere near their nests, and this is of some
importance in connection with the meaning of their courtship-
actions.
As there were thirty pairs or more on the water, I was never
at a loss for “material.” In fact, it was often very hard for my
pencil to keep up with the birds’ actions.
Some of the watching was done concealed in the boathouses,
and some from a screened punt, but the major part from the
bank. This is in many ways the most useful. With ‘good
instruments * (in this case Goerz-Trieder binoculars x12 and a
telescope x30) every action can be easily followed, the birds
are not seared, the field of view is uninterrupted, and it is far
easier to follow the actions of the same pair of birds for a long
period of time.
This, as I say, is best for discovering the general course of
events; but just as the microscopist must for certain details
supplement his low-power lens with an immersion objective,
so here, watching at close quarters must be adopted in order
to work out the exact meaning of each separate bit of behaviour.
Only when the general course of events has been roughly traced
and some hypothesis, however vague, framed concerning it in
the watcher’s mind, can the fine shades of behaviour have any
meaning for him. It is impossible to notice or record everything,
and only when some general idea has been gained can the value
of any fact be properly appreciated. It is on this account that
IT would say, always begin by distant watching; otherwise you
will not be able to see the wood for the trees.
My brother, Mr. N. T. Huxley, spent much time watching,
and several of the incidents here recorded are from his notes.
His help was most valuable, and I wish to acknowledge it here.
To Mr. James Street, Head Keeper at the Tring Reservoirs, I
am indebted for much information, and for his help in arranging
hiding-places, ete.
8. ABSTRACT OF SELOUS’S WoRK.
T venture to append a short abstract which I had to make of
Selous’s diary notes for my own use, in hopes that others may find
it useful too.
* A simple apparatus, which makes the task of simultaneous note-taking and
observation very much easier, may be constructed as follows :—On to a folding
camera tripod is screwed a ball-and-socket camera-holder (special telescope-holders
can, I believe, be purchased) ; the field-glasses are clamped, by means of a long
screw and nut, between two leather-lined pieces of wood, and the lower piece of
wood can be screwed on to the platform of the ball-and-socket. Both for stationary
and moving objects the fatigue of observation is enormously lessened hy this means ;
in addition, one or both hands are left free, and so notes can be taken while
watching—a necessity, almost, for reliable work.
330 MR. J. S. HUXLEY ON THE
The place of observation was a single large sheet of water.
In 1900 he watched, fairly continuously, from April 27th to
May 25th, except for a break of nine days (May 8th to 17th).
At first there was but a single pair of Grebes. A nest was built,
probably shortly before May 3rd. On May 3rd the birds paired
(or attempted to pair) on the nest. A single egg was laid by
May 38rd, a second before May 8th; then came a gap, and on
the 17th the nest was destroyed bya boy. There was nowa third
bird, an odd male, on the lake. The hen ‘‘flirted” with him, but
the “right” cock drove him away: and, although he stayed till
the 22nd, he apparently remained alone and disconsolate all the
time. In a single day (May 20th) the pair built a good portion
of a second nest. This they continued to build during the 21st,
the cock meanwhile building (by himself) a rudimentary nest or
platform close to the bank. On the 22nd they were building
another (third) nest in the reeds, the cock again building a
(second) platform by the bank. On the 23rd there was an
unsuccessful, and later a successful, attempt at pairing on the
nest. On the 28th the nest seemed abandoned ; however, when
Selous returned, towards the end of July, there were two nearly
full-grown young.
In 1901 he watched for a good part of April, then nearly
continuously from April 22nd to May 14th, and then off and
on till nearly the end of May. There was again a single pair
on the water, and he believes these to have been the same birds
as were there the year before. Before he started watching
they had made a kind of nest, but a very poor one—a mere
sodden heap of weeds scarcely showing above the surface and
not at first sight to be easily distinguished from the growing
weeds about it. ‘This appears to have been only a pairing-
platform. On the 25th they paired on this platform. On
May 2nd they executed a regular weed-trick and Penguin-dance,
and, some time afterwards, paired again on the platform. As
time went on they grew less and less interesting, and it finally
grew clear that they were not going to lay. On June 12th,
when Selous visited the place after a fortnight’s absence, there
was no sign of the birds—they had gone for good.
That is the bald diary ; now for the birds’ behaviour. Under
different headings I will summarize the actions of the ‘1900
pair,” and the ‘1901 pair.” (In passing, be it remarked that
Selous has no proof that the birds he saw in 1901 were really
the same as those of 1900. He says:—“‘ As they were the one
and only pair on the same sheet of water, and as the nest was
in approximately the same place, I assume and feel personally
quite certain that they were.” However, there are certain
definite differences in behaviour in 1900 and 1901, which make
it at least possible that they were not the same pair.)
Nest-building.
(2) 1900.—Both birds may help in building the nest, usually
diving to fetch weeds from the bottom, but sometimes gathering
COURTSHIP OF THE GREAT CRESTED GREBE, Ns)
them from the surface. The mass of weeds brought up by the
bird may be very large—a good deal larger than the bird’s head,
indeed, with streamers trailing beyond the tail. One bird
(usually the male) might bring weed to the nest while the
other was incubating; when this was so, the sitting bird would
generally arrange the weeds with its beak, though sometimes
both would arrange the weed together. Weed was added to the
nest for at least five days after the first egg was laid.
In the building of the second nest, he saw one morning the
cock build a few minutes alone; then both cock and hen build
together very hard for about forty-five minutes (74 cargoes in
forty minutes); then the hen build a little by herself. The cock,
meanwhile, after a short rest, began building a platform, acting
precisely as when building the true nest. After fetching 28 cargoes
he stopped and rested. The next morning the same great activity
was visible; but now the birds were building a third (true) nest.
In fifty minutes (including a'pause) they brought 100 cargoes of
weed, the last 10 or so being brought by the cock alone. This
time the cock did not at first desert the nest altogether in favour
of his platform, but every now and then diverted a cargo of weed
to his own private platform-use. After a rest, however, he
reversed his former behaviour; he now began working syste-
matically at the platform, but occasionally took a cargo to the
nest. Sometimes he seemed to hesitate between the nest and
the platform. The next day there was a little more building,
mainly by the cock, and after this no more records.
As to the part played in nest-building by cock and hen
respectively, Selous says in regard to this pair :—‘‘ The interest
taken by the male in the nest has been very marked throughout,
more so even—in appearance, at any rate—than that of the
female, though in the actual building of it she has been yet
more efficient than he” (/.c. p. 179). Although he never carried
quite so large a cargo as the hen sometimes did, yet his average
was as good as hers, and when he swam with his burden to the
nest he went much faster.
Sometimes the cock would pass his cargo of weeds to the hen,
who (if she did not drop it) would put it on the nest. He never
saw this action reversed, nor did he even see the heu help in the
building of the cock’s platform, or building one for herself alone.
The hen alone brought large sticks to the nest (however, the cock
was seen to bring a stick to his platform).
In the only recorded case where a nest was watched during
incubation (Selous, J. c. pp. 161-170), the cock alone brought
weeds to the nest, though the hen might arrange what he
brought. This is probably of no importance. (The bulk of the
nest was presumably built beforehand by both birds together.)
(6) 1901.—This year the only “nest” seen by Selous seems to
have been a mere pairing-platform, the actual building of which
was not observed. Occasionally the birds would add-to it, but
in a very perfunctory-looking way, and never more than a few
bits of weed at a time,
532 MR. J. S. HUXLEY ON THE
Courtship- Actions.
Selous observed numerous bouts of shaking, which he refers to
in various ways: ¢.g., ‘They front each other in the water, and,
with their snaky necks reared up, ¢dter a little with the beak, or
make little tosses of their heads in the air” (loc. cit. p. 341).
He has not, however, attended accurately to the positions of ruff
and ears. All that can be said is that his pair (or two pairs)
of birds certainly went through the ceremony of shaking, and
apparently in just the same way as the many pairs seen by me.
As far as I can judge (though judging is difficult) they did not
shake quite so often. The important thing to notice, however,
is that they did shake, even when they were quite alone on the
water. That either jealousy or choice of mates should be the
immediate cause, or purpose, of the action is thus absolutely
excluded.
On p. 457 he says:—‘‘They front each other with reared necks
in the way often alluded to; then, without tdter-ing, each throws
up the head several times into the air, at the same time opening
and closing the long, slender bill.” He obviously considers this
as being different from the usual ceremony, and adds that he
has seen the same action several times, though less pronounced.
I think it probable, if not certain, that his eye was here simply
caught by a somewhat more pronounced shake than usual, the
process referred to as tdéer-ing being then what I should call a
bout of languid shaking; but in the absence of further details
one cannot be sure.
A possible but rather rudimentary display ceremony is perhaps
indicated on Selous’s p. 340:—‘Once, too, the male flies suddenly
some way off over the water.”
Then on p. 343 is described a very fine weed-trick and Penguin-
dance. It started with a bout of shaking; then the hen dived
and came up with a small piece of weed which she apparently
dropped. Just before or just after the hen came up (probably
before, to judge from my experience), the cock dived too, and
brought up a large bunch of weed. They came face to face,
and “all at once both leaped entirely upright in the water.”
The hen took hold of the dangling end of the weed which the
cock was carrying, and then they “chasséd,” “with little waddling
steps” from side to side (in the case seen by me, the birds rotated
slightly back and fro on their axis and did not actually move
from side to side. I think Selous is mistaken: such an action
as he describes would be impossible on open water). Finally
they sank down again, the weed was dropped, and “the male sets
off, full of intention, to the nest on the opposite shore.” After
some time the act of pairing was gone through. This is im-
portant as showing that this elaborate courtship-action may
sometimes lead more or less directly to pairing, i.e. may act
(more or less) as an excitant.
Finally, I must just refer to two more scenes. (1) (p. 163) :—
COURTSHIP OF THE GREAT CRESTED GREBE. 533
*¢ When just in front of each other one dives and brings up some
weed, which they both discuss in the friendliest manner, pulling
it about, and perhaps eating a little.” (2) (p. 340) :—‘ Once one
of them—I think the male—comes up with something in his bill,
which he dabbles about on the surface and seems to sport with,
the other coming close up and appearing to take an interest.”
This something Selous thinks was a bunch of weed. These
actions may bear some relation to the weed-trick, and at all
events, even if the main purpose was feeding, the common par-
ticipation of the two birds denotes that some sexual flavour
attached to the act. More light is needed on the habit. (It is,
perhaps, connected with the arranging of the weeds on the nest
by both birds together, as described on Selous’s p. 162.)
Relations of the pair with other birds of the same species.
For some days a solitary male appeared on the water where
the single pair was living. The hen of the pair apparently *
indulged in a little flirtation with the odd cock—a bout of
shaking. This roused the jealousy of the rightful husband, who
approached in the usual threatening ‘ Dundreary ” attitude, and
dived to attack the third bird from below the surface, repeating
the diving attack a second time. Later the rightful couple were
together and apparently bore. down purposefully upon the odd
male. The hen rested, while the cock drove his rival away
by the attacking dive, and then returned, to go through an
“excited” bout of shaking with his mate; first, however, “he
swims about for a little, with the head still lowered, and in a
proud sort of way.”
Thus, as far as jealousy is concerned, Selous’s observations are
in agreement with mine.
Pairing-Actions.
The most important of Selous’s observations are concerned
with nest-building and pairing; indeed, the full sequence of the
pairing-actions seems to have been witnessed by his eyes alone.
I have already given a general account of his observations and
the deductions to be drawn from them. Here I have simply
tabulated some of his detailed descriptions.
1900.
(a) One bird on the nest.
(1) May 3rd. The hen was on the nest; the cock swam up and
attempted to pair. There is no record of the hen previously
“going” into the passive position, but as this was at the
very beginning of Selous’s observations, he may well not
have grasped its significance.
* T say apparently, for Selous is not quite certain as to the birds’ identity, This
description, however, agrees excellently with my observations.
534 MR. J. S. HUXLEY ON THE
(2) May 3rd. After this attempt the hen continued to sit ; the
cock returned at intervals, and at one of his returns Selous
noticed the hen assume the passive position. (Here there is
again, perhaps, an error of omission. See the next entry.)
(3) May 4th. The hen was sitting; the cock approached at least
seven times, and at each approach the hen went into the
passive position. In these last two cases (2 and 3) the cock
paid no particular attention to the hen, save for the mere
fact of his approaching the nest.
(b) Both birds close to the nest.
(4) May 21st. The birds were resting after having built most of
a second true nest; they then began building again, and
after about a quarter of an hour the hen jumped up on to
the nest and assumed the passive position. The cock made
no response. She soon came off, and the building went on.
(5) May 22nd. Almost exactly the same scene as (4) on the
previous day.
(c) Both birds approach the nest together from a distance.
(6) May 23rd. (Not recorded which bird led the way.) The
hen assumed the passive attitude on the water; when she
stopped the cock did the same, but remained in the attitude
longer.
(7) May 23rd. After (6) there was a pause of about forty
minutes, during some of which the cock (alone) added to
the nest; he then ascended the nest and assumed the
passive attitude. The hen came up several times, and each |
time acted as if about to leap up into the active attitude.
Meanwhile the cock rose once or twice and then sank down
again into the passive attitude. Finally he gave it up and
took to the water.
(8) May 23rd. After (7) there was a pause of a few minutes,
during which the birds separated and went quite far afield.
Then the hen ascended the nest and assumed the passive
attitude. The cock came up, behaved just as the hen had
done before (in 7), but finally leaped up, and there was an
attempt to pair.
[I have put (6), (7) and (8) together because each one seems
to lead up to the next. If we were to separate them strictly,
(7) should be under heading (b), for the birds remained fairly
close to the nest all the time between (6) and (7). |
(9) May 23rd. Nearly an hour later. This has been already
described (p. 503). The hen incited the cock by lying along
the water ; the cock responded by also going into the passive
attitude ; the hen ascended the nest and assumed the passive
attitude, and the cock then attempted to pair.
Thus, in 1900, the three actual attempts at pairing were made
COURTSHIP OF THE GREAT CRESTED GREBE. 535
by the cock. Of the “ incitations to pair” (when one bird goes
into the passive attitude), seven were made by the hen and one
by the cock. In the case of this one, the cock ascended the nest
directly ; as to the hen, on two occasions she was already on the
nest, on three she ascended the nest directly, and on two she
assumed the attitude on the open water.
1901. (Only a pairing-platform available.)
(a) One bird sitting. (No instances, because the birds apparently
do not sit on the pairing-platform.)
(b) Both birds near the platform. No record. (There are two
doubtful records where he first notices the birds already at
the bed of weeds where the platform is, but here probably
he had simply not noticed their previous approach.)
(c) Both birds approach the platform together from a distance.
(Fifteen records, counting the two doubtful ones.)
(i.) One bird swims straight to the platform, ascends, and
assumes the passive attitude.
This was seen ten times ; once it was done by the hen, the
other nine times by the cock.
(ii.) One bird assumes the passive attitude on the water near
the platform.
This was observed five times; twice it was the female, once the
male, and twice the sex was doubtful. (Here, therefore, there
is not the preponderance of incitations by the male that was
seen in (i.)).
On April 25th the hen assumed the passive attitude, and
nothing further happened (immediately).
On May 11th the hen went into the passive attitude, upon
which the cock followed suit by going into the attitude too.
Some time afterwards the scene was repeated, but with the parts
played by the sexes reversed (cf. 1900 (6) & (9), above).
This passive attitude, adopted successively by both birds on
the water, might or might not lead to one of them ascending
the platform and there assuming the passive attitude.
Very often an incitation might have no immediate result,
but after a short pause further pairing-actions might be gone
through. I will quote a couple of Selous’s records for whole
mornings.
April 25th.
(a) (There have already been several approaches to the neigh-
bourhood of the nest, and several bouts of shaking.)
The hen goes into the passive attitude near the nest ;
but there is no result, and both swim away.
(3) “* Very soon afterwards” they return, and the cock goes
straight to the platform, where he assumes the passive
536 MR. J. S. HUXLEY ON THE
attitude. The hen very shortly jumps up, and pairing
takes place.
(y) A little later they again approach the platform, and the
cock again goes into the passive attitude upon it.
The hen, however, takes no notice, and the cock
comes off.
(8) He follows her, they both turn, and he repeats his
previous action—again without result. He then comes
off, and fetching a piece of weed, lays it on the nest.
The female comes up, and they lay a few bits of weed
on the nest together, but very perfunctorily.
May \\th.
(a) Not long after a bout of shaking, they swim together to
the weeds. The hen assumes the passive attitude on
the water. The cock approaches, “‘ appearing interested”
(cf. p. 501), but suddenly turns round and also assumes
the passive position (but not so pronouncedly), in such
a way that the two are tail to tail. Both then rise
up, the cock presses past the hen, and goes into the
passive position on the nest. The hen makes but a
slight response, and the cock, after adding a piece of
weed to the nest, swims off in company with his mate.
(8) After less than half an hour they swim towards the
platform, the cock leading. The cock goes into the
passive attitude, and is imitated by the hen when she
arrives. There is, however, no further result, and
the pair swim off in company.
(y) After about forty minutes they again swim towards the
platform; the cock is far ahead, and on reaching
the platform he ascends it and assumes the passive
attitude. On this, however, the hen apparently
becomes coy, for she suddenly turns and swims off;
but when the cock follows her, she turns and swims
eagerly to him—a pretty piece of psychology.
With this I have summarised the most important of Selous’s
facts. Others will be found under the separate headings.
9. FURTHER DETAILS REGARDING THE RELATIONS OF THE SEXES.
(i.) Shaking.
The typical bout of shaking is of ten or a dozen shakes, the
crest erected in a definite way, the necks stretched straight up to
their fullest extent, the two birds facing each other at a distance
of a few inches only.
It is usually initiated by the birds swimming towards each other
at a moderate pace, meanwhile gradually raising their crests and
necks, and giving a repeated double call rather resembling that
up-and-down call of the Snipe as he sits in the marsh. i
COURTSHIP OF THE GREAT CRESTED GREBE. 537
At the beginning of a bout the attitude often seems to express an
extra degree of excitement, but especially so after a “flirtation”;
the ruff is a little more circular, the ears pressed forward to their
limit, and the neck curved over a little at the top, so that the
heads and beaks are pointing somewhat downwards; this attitude
never lasts long, and soon subsides into the ordinary one, in
which the birds give somewhat the alert impression of a couple
of smooth-haired Fox-terriers. Asa general rule, habit-preening
does not begin until after a few shakes, and usually gets a little
more frequent as the bout goes on.
Usually, the bout is closed by the two birds simply drifting
slowly apart, and gradually lowering their crests, or else one or
both of them may turn sharp up into the wind with a more
sudden closure of the crest.
In the longest bout seen the pair gave 84 shakes, while in the
shortest seen (which is also the shortest possible) each bird only
gave a single shake.
Often there are somewhat abortive bouts, without enthusiasm
on the part of either bird; these do not last long, never for more
than seven or eight shakes, and the crests and necks are often
not erected to their proper position. There are degrees of excite-
ment. In the lowest the neck is in the graceful curve of the
ordinary swimming position, the ruff is relaxed, and the ears are
searcely half-raised. In the next stage, the ears are fully raised :
then, the ruffs are slightly expanded as well: then, the neck is
raised more and more: and finally both ruff and neck come to
their typical extension.
A certain frame of mind is necessary for shaking, and some-
times even the expressed desire of the other bird to shake cannot
arouse this state (see p. 544); for still further details the reader
is referred to section 10.
(ii.) Nest building.
There is one curious habit connected with nest-building about
the significance of which I am not at all sure: this is the tram-
pling down of the nest. Lying in a punt in the reeds I have
heard this trampling, first on one side, then on the other—
squelch, squelch, squelch,—sounds of some creature trampling
heavily with alternate feet on something sodden.
This is the water-birds treading down their nests. The Grebe
does it, and, I believe, the Coot, and possibly other birds as well.
The one occasion where I saw it well is worth recording, partly
on this account and partly for another reason, as will shortly be
seen. A pair had been fishing and resting; the cock then stayed
perfectly still, not very far from the nest, for some minutes, and
the hen went off and fished. The cock moved slowly towards
the nest, and was there joined by the hen; after a short time
the hen got on to the nest, there, in the usual ungainly upright
position, stamped heavily twelve or fourteen times on the sodden
weeds, and then settled down into a sitting position She
538 MR. J. S. HUXLEY ON THE
remained thus for about two minutes, then got off, apparently
put a few bits of weed on the nest, and swam “off with the cock,
who for his part had remained quite quiet all the time. When
I went to inspect the nest later in the day, I found that there
were no eggs, and that it seemed to be a mere pairing-platform—
old, sodden, low, and covered with excreta.
This is thus the only case on record where a bird has ascended
the pairing-platform or nest except for the purpose of incubation
or to go into the passive pairing-attitude.
The cock of course uses his special platform to rest on, so it is
possible that the pairing-platform is used by the hen as a corre-
sponding resting-place. Or, in amplification of what I have
already suggested (p. 518), that when the true nest is built the hen
incubates longer and so uses that as a resting-place, the cock has
his platform, and both sexes use the pairing-platform.
A third possibility (but not a very likely one) is that the
sitting on the nest (platform ) in this case was in reality only
the first step towards assuming the passive attitude, but that the
bird’s sexual excitement was not high enough to ‘complete the
action.
Most probably this, like many other bits of behaviour, was an
“accidental” and useless release of ener ey, rendered possible by
the mechanism of the bird’s mind.
(iii.) Details of the Relations of Different Pairs
to each other.
I have thought it worth while to go into this at some length,
in order to show how extremely complicated the birds’ mental
states are, and how like may at a moment’s notice be turned into
dislike. We will consider the relations of a pair with an odd
bird in the neighbourhood.
Let us for brevity’s sake call the three birds X, Y, and “ Y.”
“VY” is the intruder, whom we also call the odd bird; X and Y
are the pair, or the paired birds, cock and hen; Y is of the same
sex as “ Y,” X of the opposite sex. This will serve when we
want general formule. In particular cases, where the sexes
have been accurately observed, we can employ a similar formula,
eg. 6, 9,“9, or 6, 9, “6d,” according as the odd bird is a
cock or hen; or we can simply say “Y”=“ 3” or “9” asithe
case may be.
Now for our general statements :—
1, The disposition of X towards ““ Y” may be (a) well-disposed
—in other words, X may be at the moment flirtatiously
inclined in general.
(6) Indifferent. Then X does nothing in particular, and Y
drives “ Y” away.
(c) Hostile. Then X helps Y drive “ Y” away ;
but the actual initiative, the first step towards a “flirtation,”
may be taken (1.) by X, (i1.) by ‘“ Y.”
COURTSHIP OF THE GREAT CRESTED GREBE. 539
2. The disposition of Y towards “ Y”; this is always more or
less hostile, but there are variations (w) in the time at which the
hostility is first shown, and (4) in the way in which it is shown.
(a) Time. Y becomes hostile
(i.) on passing near ‘‘Y,” although “‘Y” has been quite
inactive ;
(ii.) on passing near “ Y,” but only after ‘‘ Y ” has first gone
into the hostile Dundreary attitude ;
(111.) only when X (its mate) makes as if to approach “ Y” ;
(iv.) only when its mate actually begins to shake with ** Y” ;
(v.) only after its mate has shaken for some time with “ Y.”
(b) Method.
Y may (i.) simply swim at “Y”;
(i1.) assume the hostile attitude and swim towards “ Y” ;
(i11.) fly at ‘“ Y” along the surface of the water ;
(iv.) dive and endeavour to come at ‘‘ Y” with the beak
from below the surface.
In addition, these actions may be gone through in succession ;
if so, they are always gone through in this order, except that (iii.)
and (iv.) may be reversed. ;
3. The disposition of ‘‘ Y.”
““Y” may be (a) simply indifferent to the presence of the pair ;
(6) hostile to the pair ;
(c) eager to “ flirt” with X.
Here again the initiative may be taken (i.) by X, (ii.) by ‘“ Y.”
Combinations of these pretty well exhaust the possibilities ;
here I shall give an idea of the most usual happenings.
When the two birds of a pair are swimming along together,
and they pass close to a third bird, X usually takes no notice,
and Y goes into the Dundreary attitude while passing “ Y,”
sometimes swimming a little towards ‘“Y.” The odd bird, “ Y,”
may be, and usually is, entirely indifferent to the pair, though it
may adopt a threatening (Dundreary) attitude as a response to
the similar threat of Y; and I once saw ‘“* Y” take the initiative
in threatening.
On the other hand, I have never seen any sign of a flirtation
between X and the odd bird in these circumstances.
That X should be willing or desirous to flirt with “ Y,” it
seems necessary that its mate should either be absent or fairly
distant, or, if close at hand, lethargic (see section 10, record 1), or
unwilling to perform any courtship-actions. When a. flirtation
does ensue, “ Y” may have taken the initiative, by swimming
close up, calling, or going into the searching Dundreary attitude ;
or X may have taken the initiative by swimming up te a
Proc. Zoot. Soc.—1914, No, XX XVIII, 38
54O MR. J. S. HUXLEY ON THE
perfectly innocent “Y.” The former, from my records, seems to be
more usual (as one would expect: it is more probable that a lone
bird will be more eager to “shake” than one whose mate is near
at hand).
One interesting fact emerges from table A (p. 542): in seven
out of eight cases where there was a flirtation and I could be sure
of the sexes, “ Y” was a female, while of the cases where X was
indifferent or hostile to “ Y,” ““ Y” was female in about half the
number.
This is probably not merely chance; I believe that here the
male Grebe possesses a little more of the normal characteristics
of males.
I have several records where “ Y” is very threatening from
the first; here the pair always seem to make common cause
against the intruder. I do not see how these cases can have
anything to do with a desire of “ Y” to shake, ete., nor is there in
the behaviour of the pair any sign of jealousy. ‘There seems to
be only hostility, and [am disposed to think that in all such
eases the pair has been trespassing on another's territory
(see p. 558).
Y may drive “ Y” right off, or content itself with going into
the threatening attitude. ‘“Y” is always driven off if Y sees a
flirtation going on; but when the pair are simply swimming past
the odd bird, Y contents itself with going into the threatening
attitude (in one doubtful case only was “ Y” driven away) ;
here we have association at work. In cases where “ Y” hangs
about, it may be driven off by Y even though no flirtation takes
lace.
If Y wishes to drive “ Y” off, it usually dives. Sometimes
““Y” is completely taken by surprise, and, as it flies off, its place
is at once taken by the jealous one. Presumably Y’s beak
sometimes actually comes into play, which must be very. un-
pleasant for ‘“Y.” At other times “‘ Y” sees Y dive, and is off
at once.
The mere presence of an odd bird is not sufficient stimulus to
induce a bout of shaking between the pair. Shaking is usual
(though not invariable) after a flirtation and subsequent driving
away of ‘ Y.”
When a bout of shaking does take place after flirtation, it
always seems to begin in the forward position. This position
in its full development I have never seen except after an odd
bird has been driven away, so that we have here an interesting
example of a definite form of courtship-action used exclusively
under the influence of jealousy.
Let us close with one or two interesting cases—actual hap-
penings. Here is one:—
1. The cock of a pair, saw an odd bird (? sex) near by, and
drove it off, by flying at it, spluttering along the water. He
then came back towards his mate, and from his attitude I thought
COURTSHIP OF THE GREAT CRESTED GREBE. 541
he was going to shake with her. However, he then saw another
odd bird (¢ sex) not far off, and, his pecker presumably being up,
went off and drove it away too. After this he came back, and
a short but vigorous bout of shaking ensued. I do not suppose
he would have driven the second bird off if he had not been roused
by the first.
2. Then a second:—An odd cock was seen ‘in a very threat-
ening attitude,” some twenty yards away from a pair. They got
close together, the cock going into a fairly good threatening
attitude: they then swam, the cock leading, towards the intruder,
but suddenly turned tail; however, they soon faced round again,
and waited a bit. ‘ Y” was now only about ten yards off. ‘Then
the pair swam a little away (this is very odd) and then all three
dived ; asa result, ““Y” was driven a short distance away, and
the pair made as if to shake, but did not. “ Y” approached
again, and the same scene was re-enacted almost identically (most
ludicrous to watch, it was !); finally came several long dives on
the part of all three birds, and “ Y” was driven right off. Strange
to say, no shaking followed.
Here, in face of an obviously hostile third bird, the pair
united at once in common action (cf. Selous, in the case of Ring-
Dotterel).
3. Once I saw the odd bird approach “like a dolphin ”—pro-
gressing for the most part subaqueously, but now and then lifting
first head and then back out of the water, only to disappear
again. What this may mean I do not know.
4, In one case where there had been a flirtatious shake (‘ Y ”
= 2), the rightful hen came up, and instead of at once driving
“Y” away, started shaking, so that for a short time there was a
parti a trois. She then drove “ Y” off, and then returned and
had a long shake with X. This I have only seen once.
5. Once where “ Y” (a hen) had called, X swam almost up to
her, but at the last moment was seized with a ‘fit of repentance,”
returned to his mate, and shook with her. Perhaps it was not
repentance at all; perhaps on seeing her mate go off towards
“Y,”’ the hen gave some sign that she was ready to shake, or
rather that she was roused enough to shake. This would be all
the cock wanted, and, seeing this, he came back. This is only a
possible explanation, but it at least has its parallels in our own
affairs.
6. A variation on the above was given by the same cock a little
earlier; this time he had actually shaken a bit with ““ Y”, but on
seeing his mate approaching, he suddenly turned on “Y” and
drove her off. The rightful hen did not trouble herself further ;
but there was, curiously enough, no shaking on the part of the
pair.
7. An odd bird approached; Y (? sex) swam towards it. Both
went into the threatening attitude and remained perfectly
motionless, looking very fierce at each other at a distance of only
38*
542, MR. J. S. HUXLEY ON THE
five or six feet, calling (‘‘ barking”) the while. - At length Y
came up with a loud trumpet call, there was a great flying of all
three, “ Y” was driven off, and the pai had a short bout of
shaking.
This is something like 2—hostility evinced from the start by
“VY,” the pair acting together against the intruder,
8. I only once saw two pairs come into conflict; and un-
fortunately could not make out much. There was much diving,
and, finally, two birds went off together; the other two had a
short bout of shaking.
Finally come the two cases where apparently two “odd birds”
meet.
9. In one I saw a short bout of shaking. Then the cock went
off in the search (Dundreary) attitude; the hen, though quite
close, took no notice, and the cock finally went right off. I am
almost sure it was this same hen who later shook with another
cock.
10. Jn the other (p. 546), a cock came flying over, settled near
a hen, and they had a short bout of shaking. Then the cock dived ;
the hen still kept her ruff up expectantly, but the cock came up a
long way off, swimming away from her, and she put it down ;
and so the scene ended.
In both these cases it is pretty clear that the birds were not a
pair, but that, finding themselves together, they tried a bout of
shaking. This, however, was somehow not satisfactory—it was
not what they were accustomed to; and they parted. Both
times one bird (as it happened, the male) was obviously searching
for his mate, and it may be that this “ prepossession’’ led to the
flirtation being quickly broken off.
(RABEE AL
Disposition of X to “ Y.”
AS Les = Ey
Sex of SiYer
| lie = erase a H
3 2 P
| (@) X flirtatiously disposed to “Y” .........-........ sy ie
(OD) PPXGindifterentytopVacu se oe 4 | 3 2
(Oxo rostilertow ayy et Ue a iS) Ste ik ila we 1 | 2 2
* In two of the seven cases, X (3) later became hostile to “Y” when his own
mate approached, and in a third case he was almost entirely indifferent to “ Y.”
COURTSHIP OF THE GREAT CRESTED GREBE, 543
TasLe B.
Disposition of ‘‘ Y ” to the pair.
Ficures in brackets include doubtful possibilities.
|
_A,—X and Y (the pair) swimming together in a |
definite direction.
SexOh saver
(a) “Y” (the odd bird) threatening :
(Ge) ee acenba kesh taentni tiativicmea sy eameseeeeeeece = |
(ii.) ““ Y” does not take the initiative
(Qe NE HAN CTEHODISS Hrgtoppaoceadbnes rosa onusecHnecesea||
(c) “Y indifferent ......
| B.—X and Y fishing, resting, ete.—not swimming.
|
|
vice to any future observers.
(a) “Y” threatening :
(G5) erantalkespulelimibiaytivieneessceerreerte- eer |
(ii.) ‘““ Y” does not take the initiative ....,....
(ii.) Doubtful
(b) oy flurtatious 8
(Go) peeveretakeshihepmniibiatiyer spear eee ae
(u.) “ Y” does not take the initiative
(it, Domo cacccooce
(c) “¥ ” indifferent ......
Sk Caen
1
1 |
1 5 |
1 (2) 0) |
loay|
Day Ps l1@)
2 |
1(2)| 2 )1@2) |
2 bea
10. Recorps rrom my Notes.
Here follow a number of actual incidents which may be of ser-
Among other things, they show
very clearly the individual differences between different pairs.
I have numbered these scenes, and append here a little index
showing where descriptions of various courtship activities may be
found :—
Courtship-action.
Scene.
Bout of shaking
Display ceremony ...... seinen te ceas a
Discovery ceremony
Diving alone
Weed-trick
Complete Penguin-dance ............ are
Flirtation
Cr i i i a i ii aca
ee ee ee
ee eC i a ie id
weer etree ores
Hostility to birds of other pairs
per oee!
Lo
joel
Or
Ome ams
a
~
2
~
ay
~
bs
—
Or
[xis Ja
|
ware}
Or
~
Dayo wwe pe
— ee bh bd ©
—
~~
jA44 MR. J. S. HUXLEY ON THE
1. April 10. 4.8—4.40 p.m.
A pair, ¢ and Q easily distinguishable.
When I first saw them they were indulging in a typical shake
of moderate length, ended by one turning away from the other.
After this for about 17 minutes they moved slowly in one direc-
tion, the hen always leading the way. When not swimming she
did some fairly vigorous preening. Sometimes she was 30 or 40
yards ahead of her mate, but malic she got as far away as this,
she always swam back and joined the ‘cock. (Pleasure merely
in each other’s presence, and dislike of being separated, 1
marked in many monogamous birds.) He spent most of ae
time with his head under his wing, but now and then woke
up, looked about him, and gave some rapid strokes towards
the hen ; occasionally he did a ‘Tittle preening. Not only was the
hen more active and awake than the cock; she was also more
emotionally inclined. She kept on coming close up to him and
shaking her head slightly, trying obviously to stimulate him to
respond. The first time she did this (4. 13) the cock just raised
his head from under his wing, gave a couple of scarcely-visible
shakes, without extending his neck, and relapsed into somnolence ;
while to her later advances he responded not at all. She was
very restless ; would swim up, give two or three shakes, swim a
few yards off, turn, come back, swim off again, and so on, maybe
three or four times in quick succession, then she would make up
her mind and swim steadily off, only to come back again after a
few minutes’ interval. This she did four times. By 4.26, after
17 minutes of this, she began to think of feeding, for she dived
twice. Her previous emotional state had, however, not quite
died down, for she then came back right up to the cock, though
this time without any actual shaking. At 4.32 she went right
off, and began fishing a good eighty yards away. At 4.35 she
caught a big fish, swallowed it, and went on diving. The cock
meanwhile rested and preened himself, and at 4.40 I lost sight
of both.
This well illustrates the way in which the physiological and
emotional states of individual birds vary from hour to hour.
That the male was capable of normal excitement is shown by his
shaking in the usual way at 4.8. This exhausted his emotional
fires fos the time being, but left the hen still with a good deal of
pent- up excitement. It seems (as one would expect) to be “no
fun” to shake all by oneself, and so her potential energy had to
be released through other channels, giving as a result the quarter
of an hour’s restlessness.
At 4.43 a bird which is recorded as “ 3, probably the same as
that lost sight of at 4.40,” went into the regular Cat-position,
and its mate appeared in the usual way, rising erect closely from
below the surface. (Discovery Ceremony.) If they were the
same pair, it is obvious that the half-hour’s rest had restored to
the cock all his emotional energy, and the variation in emotional
states is still more clearly brought out.
COURTSHIP OF THE GREAT CRESTED GREB®. = FAD
2. April11. 1.50—2.30 p.m.
A pair: ¢ and Q rather hard to distinguish.
1.50. I just saw the end of a shake.
1.51-2.0. They swam about vaguely, occasionally diving.
2.1. I was watching one (sex ?) when suddenly the other came
into the field of view, carrying in its beak a big bunch
(bigger a good deal than its own head) of dark, ribbony
weed, which must have just been fetched from the bottom.
The bird was swimming fast and rather low, in the ordinary
position adopted when approaching its mate with weeds. It
came right up to its mate, and the pair shook (without
habit-preening) for 10 seconds or so. Then (I am practically,
but not absolutely, sure) the weedless bird took some of the
weed, and shaking began again. This lasted a still shorter
time—“ then ” (I quote from my notes) “ both birds turned
head to wind—and lo, their ruffs were down, and there was
no weed in their mouths!” They then swam off together.
2.3-2.10. Lost to sight behind reeds.
2.11. Out again. They turned to face each other, and then
shook five or six times. To start with, strange to say, their
necks were right down in the normal swimming position.
As they shook, they gradually raised them till they were
half pear-shaped. They then stayed motionless for about
20 seconds, then shook twice, and swam slowly apart.
2.12-2.27. For fifteen minutes, as near as may be, they did
absolutely nothing—merely drifting and swimming aimlessly
about.
2.27. One preened itself ; and then they faced each other, shook
7 or 8 times, turned up wind, and swam off into the reeds.
3. April1ll. 5 p.m.
I caught a pair in the middle of a bout of shaking. There
were 7 or 8 shakes, with an occasional habit-preen, and then they
swam apart, but with their necks still straight up and crests
erected. One stayed nearly stationary; when the other had got
some fifteen yards away, the stationary one dived. It came up
close to the other, and shaking began again, much as before.
After seven shakes they stopped and went off together, only
gradually letting necks and crests subside to their ordinary
positions. It must have then been feeding-time, for they took
three long dives across to ‘‘ Fish Corner” and began fishing.
This scene is unusual, for diving as a part of courtship-ritual
is usually associated either with the cat-position or with the
weed-fetching. The slow subsidence of neck and crest after
shaking is also not common.
4. April12. 8.20a.m. A pair.
After drifting about for 5 minutes or so, they began shaking.
They shook 10 or 12 times, with habit-preening; they then put
546 MR. J. S. HUXLEY ON THE
their ruffs down, and drifted slowly apart. When they were
separated by about twenty yards the cock dived and came up
close to the hen, upon which the pair began shaking once more.
After a very few shakes they stopped, the cock put his crest down
and swam off at a moderate pace ; the hen, however, stayed where
she was and kept her crest up. When the male was about forty
yards off, he went into the cat-position: on looking at the hen,
I saw that she had done the same, with wings fairly well arched.
The cock had at first scarcely arched his wings. but when the hen
went into position, up went his wings to the full for an instant.
It was but an instant, for then he dived; ‘‘a ripple was seen
coming quickly towards the hen along the surface of the water
(most exciting!)”; when it had nearly reached her, the
cock appeared, slowly erecting himself out of the water in the
usual way. He seemed to be facing her all the time. He
settled down, and a very long shake began. There was no
habit-preening for the first ten or a dozen shakes, nor very much
at any time. Eventually they drifted apart, put their ruffs
down, and did nothing in particular for the five minutes or so
1 went on watching.
This shows again that courtship-diving may take place apart
from the cat-position or from weed-fetching ; and also that,
although the cat-position seems usually to be employed as a
stimulus to induce a bird of the opposite sex to do the Penguin-
dive, yet the diving bird, too, may go into the same attitude
before it dives.
5. April 12. 8.50 a.m.
A cock flew over from another reservoir and alighted near a
hen. In under a quarter of a minute they had begun
shaking. They only shook seven or eight times, with habit-
preening, and then drifted apart. Soon the cock dived; the hen
kept her ruff up, but the cock had dived away from her, and
appeared a long way off. On seeing this, the hen lowered her
crest.
This seems to show that when one bird dives, and dives deep
so as to produce no ripple on the surface, the other is left in a
state of suspense which is exciting enough to make it keep its
crest up. :
Whether the two birds were a paired couple or not could
only have been proved by further watching; but I should say
that they probably were not a pair, but that their close
proximity and the absence of their real mates excited them. The
emotion found expression in the usual actions, but then the
strangeness of the hen proved unsatisfactory to the cock.
6. April 16. a.m, A pair.
I caught them shaking. After 6 or 7 shakes they separated ;
when they were some way apart, the hen went into a feeble cat-
COURTSHIP OF THE GREAT CRESTED GREBE, 547
attitude. The cock dived, and came up five or six yards off
with a fair-sized bunch of weeds in his mouth. Strange to say,
he was in something very like the normal swimming attitude,
though his ruff was fairly well erected. On seeing him the hen
to my surprise put her crest down, turned, and swam away, and
the cock could do nothing but drop his weed, lower his crest, too,
and swim after her. Nothing particular happened in the sub-
sequent five or ten minutes.
Here, when the emotional excitement reached a certain pitch,
the hen had a sudden attack of “coyness” (ef. similar behaviour
in the female Redshank, Huxley, 12,, p. 651).
7. April 16. 1.30 p.m.
I caught a pair shaking. Suddenly, and for no apparent
reason, the hen flew off, flapping along the water. 1 followed
her, but she simply settled in an ordinary attitude. However,
on looking back at the cock again, I found him engaged in
shaking with another hen. The first hen, therefore, must have
been a casual acquaintance, who departed hastily on seeing the
rightful mate coming up. The rightful pair shook 4 or 5 times
(without any habit-preening), and then on a sudden the cock
flew a few yards off, and put himself into the best cat-attitude
IT have seen. He turned round, first one way and then the
other, just as the Peacock does when in display, and then,
gradually un-arching his wings and raising his neck, swam back
to begin shaking once more with his mate. This time they shook
about ten times; habit-preening began about halfway through,
and at the same time the ruffs were half lowered. ‘Then they
both dived nearly simultaneously, and I saw them no more (they
must have made a very long dive and got into the reeds).
This is a very good example of the pure Display ceremony (see
p. 513), here induced by the extra excitement of the previous
“ flirtation.”
8. April17. 1.30 p.m. A pair.
I saw a pair shaking; they went on for a very long time (no
notes as to habit-preening), and finally one (sex ®) dived. As it
did so, I saw the other convert its crest into an “ Klizabethan
ruft”; after a few seconds it too dived. Both came up with weed
in their mouths, fairly close to each other, and the usual Penguin-
dance was gone through, followed by a short bout of shaking.
They then put their crests down, and swam off together. To
progress faster, they took three long dives, each time going under
almost simultaneously. After the third dive they came up close
to a single bird (sex ?), which at once went into the Dundreary-
attitude, Then all three dived in quick succession, and after
some time two, which I presume to have been the original pair,
came up close together, and at once began to shake, starting in
the excited forward position. After that I lost sight of them.
548 MR. J. S. HUXLEY ON THE
9. April 17. 2.25-3.5 p.m.
2.25. A pair were swimming about fairly close to each other,
resting and preening by turns.
2.28. The hen three times went into the Dundreary-attitude
with short intervals between.
2.35. After resting for some minutes, she barked five times in
succession, and relapsed into the resting position, never,
however, shutting her eyes.
2.44. One bird swam off out of sight; the other barked several
times. After a little bit the first one came back into view,
and they both began preening themselves.
2.50. Suddenly a third bird (sex ?) came swimming towards them,
and when about thirty yards off went into the Dundreary-
attitude, at the same time giving five long, loud, rolling
barks. At once both birds of the pair put themselves into
the same attitude,and faced round on the third Grebe, uttering
at the same time a series of short, low, and quickly-repeated
barks. The intruder changed its course a few points and
went off towards the reeds; directly it was out of the way,
the pair “ got up and shook.” The shaking, however, only
lasted for a short time, and they then relapsed into their
previous state of preening and swimming about.
2.55. After they had swum out of sight and back again, I saw
one of them (sex ?) go to one side in Dundreary-attitude,
calling repeatedly, and on looking further afield discovered
the reason for this in the shape of an intruding single
bird (sex ?) (probably not the previous intruder) who
was approaching in the same position and uttering the same
ery, about thirty yards away. This single bird then dived
and came up not six feet away from the other. They were
now in an attitude I had never seen before—hbest de-
scribed as the most hostile possible form of the “ Dundreary,”
differing from the typical form chiefly in that the heads
were not quite so low down. For some time they stayed
thus facing each other, still, or moving a little forwards as if
to attack, and then at once thinking better of it, and all the
time giving the low, quick bark. Finally the second bird
of the pair came up, giving a loud grinding trumpet-call, and
then all of a sudden there was a great flying of all the three at
each other, and at the end of it one bird went off, and two
(undoubtedly the original pair) were left together ; they at
once approached and shook; the shaking, however, only
went on for a short time, and then, after a few minutes’
preening, I lost them.
Here, twice over, it is obvious that the presence of a third bird
has screwed excitement to the shaking-point ; the remarkable
thing in both cases is, that the bouts of shaking thus induced
should last so short a time, whereas on another occasion (April 16,
1.30 p.m.) a similarly-induced bout ended very differently.
COURTSHIP OF THE GREAT CRESTED GREBE. 549
10) Ape lis) “p.m:
A single bird (I think a hen) was swimming about, gave the
double-trumpet twice or thrice, and then looked about. Another
bird, some forty yards off, noticed the call and turned, and they
swam quite slowly towards each other. When they had reached
each other, they began shaking, very excitedly at first. After
six or seven shakes, the hen suddenly turned straight away from
the cock, and flew or spluttered some eight yards away. She
. then put herself into a fine cat-attitude, and began turning from
side to side—all this without uttering any call. The cock watched
her thus for several seconds, and then dived and swam just below
the surface, making a ripple, and as this approached her, the hen
drew her head down ever lower on her breast. When four or five
feet off, the cock put his head and neck out—apparently to see
where he was, for he disappeared again at once. When he
finally appeared if was three feet beyond the hen, and he was
facing away from her as he “ grew out of the water” into the
customary ghostly Penguin ; he turned to face the hen as he
subsided, and finally shook with her. This bout was only a short
one, however, and after it they swam some distance apart. After
a minute or so the hen gave two double-trumpets, but then
relapsed into the state, from which the cock had never emerged,
of doing nothing in particular, and in this state some minutes
later I left them. |
This differs from the typical Discovery ceremony in two points:
(1) the birds do not usually swim together thus, but one dives at
once, (2) the cock does not generally wait and watch the hen’s
display (“ cat-attitude ”) before making his “ ghost-dive.”
11. April 18. 10.40 a.m. to 12.17 p.m. A pair.
10.40. I saw a pair close together, the cock and hen easily
distinguishable.
10.45. They preened close together for some time.
10.48. They swam off together and got close inshore.
10.52. Both dived once or twice, I think for fish.
After preening themselves for a bit (10.56-10.59) they began
diving again near the bank, and the cock caught a small fish.
11.0. They came up close together and began shaking; but they
only gave three or four shakes, and their ruffs were scarcely
half expanded.
11.2. Again after a dive they came up close together, and again
had a bout of shaking. This time they shook 10 or 12
times, and their ruffs were well up; there was no habit-
preening.
11.8. After some minutes’ swimming about further out from
the bank, occasionally picking up things from the surface of
the water, they swam together with outstretched necks
(the forward swimming - attitude), the hen swimming
much the faster. As they approached, and while their
necks were still stretched forward, they began to shake, but
MR. J. S. HUXLEY ON THE
they never put their ruffs properly up, and after five or six
shakes, during which the necks were gradually raised, but
not to their full height, the shaking degenerated into habit-
preening, and this into real preening.
11.10. They swam off and began diving again near the bank. After
the second dive they came up only about three yards apart,
and both shook their heads three or four times; the shaking
was not very vigorous, and ‘‘ had not much reference to the
other bird ”—i. e. they did not come and face each other in
the customary way. They then swam out from the bank.
As they passed a solitary hen some way off, the hen of the
pair, who was between her cock and the single bird, went
into the Dundreary-attitude. From 11.12--11.22 they swam
about, picked things off the surface, fished, and took long
“‘ progressive ” dives, ending up near the opposite bank of
the reservoir.
11.23-11.28. After a dive they came up fairly close together,
and swam towards each other with outstretched necks, which
they gradually raised as they neared each other, beginning
to shake their heads at the same time. <A prodigious bout
of shaking ensued, and was followed by diving for weed,
swimming together, beautiful penguin-actions, and final
bout of shaking. The whole thing has been already de-
scribed in detail in section 4c, p. 499. Here suffice it to say
that the hen began habit-preening before the cock, and once
begun, practised it more than he. She too dived first, came
up first, and had more weed in her mouth.
11.29-11.39. They swam back towards where I first saw them,
often picking things off the surface. A. solitary cock was
close to the line of route, and our cock went into the
Dundreary-attitude as he passed the odd bird.
11.40. They dived. The cock came up first, and gave a couple
of shakes “to himself” (cf.11.10). Then the hen came up,
and they shook together four or five times, but without
raising their crests at all, or their necks to their full extent;
they then went on fishing near the bank.
11.49. They stopped fishing; the hen began preening. The
cock swam towards her from some thirty yards off. She
came a little way to meet him, and they shook seven times
with their ruffs half-up. The bout ended in habit-
preening.
11.56. After swimming about and preening they had another
shake. This again was a very long one, like that at 11.23
(I did not count the number of shakes, as I was more intent
trying to make out various details of attitude); indeed it
was almost a precise replica of this previous long one, and
had the same sequel—a fine ‘ Penguin-dance.” The only
differences I could see were that their ruffs were not quite so
“sun-like” before diving, that both brought up plenty of
weed, that I am almost sure a good deal of the weed was
COURTSHIP OF THE GREAT CRESTED GREBE. 551
eaten, and that the final bout of shaking was less than half
as long.
12.0-12.15. The birds now swim about, preen themselves, dive,
and pick things off the surface. The cock is now more
active in searching for food than the hen, while previously
the reverse had been true. At 12.8 they passed another
bird (sex ?). My hen went close up to it, then swam rather
rapidly away, then close up again. However, nothing
happens, and neither bird goes into the Dundreary-attitude.
12.15. They pass neara single bird, which I think is a hen—the
cock, who was leading, took no notice of it; but the hen
went into the straight-necked (or angry) Dundreary-attitude
and, without giving any call, swam at the third bird. ‘The
solitary one turned, swam away, and finally flew some
fifteen yards off, upon which my hen turned, assumed
a normal attitude, and swam back towards her mate. No
shaking or other expression of emotion, however, took place.
12.17. They began fishing close inshore.
12.18. I took my eyes off them, and when I looked back could no
longer be sure of them among the several birds along the
shore.
My notes on this pair I have given in full because I had them
under continuous observation for a considerable time (over an
hour and a half). Their behaviour is of interest in various ways.
In the first place, we see how, in this pair at least, outbursts of
violent emotional actions alternate with calm periods during
which the birds rest or feed, occasionally indulging in a short
and rather languid bout of shaking. When passing near a third
bird, one of the pair usually went into the threatening attitude
(Dundreary). In every case I could be sure about, the bird that
did this was of the same sex as the third or single bird. The
pair had marked idiosyncrasies of its own, both as regards what it
did do and what it did not do: and besides this, both cock and
hen had tricks of their own in performing the courtship-actions,
which I think were certainly permanent and not due to changeable
physiological states.
All this took place on the reedless reservoir, where there are no
nests, and consequently far away from the scene of actual pairing.
Physiologically, therefore, the ‘‘ courtship ” and the act of pairing
are entirely detached from each other.
12. April 18. 2.30 p.m.
A pair shook about forty times, with habit-preening. At the end
they turned their backs on one another, still shaking and habit-
preening—to themselves, as it were,—but gradually letting their
erests sink. When about twenty yards apart they turned, swam
slowly together, and shook a little, but without raising their crests.
Then for about a quarter of an hour they stayed quietly facing
each other, preening themselves. After this there was a short
bout of shaking, then a rest, then another bout of some thirty
552 MR. J. S. HUXLEY ON THE
shakes. This, like the long first bout, was followed by their turning
their backs on each other and shaking “ to themselves ”—a couple
of shakes and 6 or 7 “ habit-preens.” This time, however, instead
of turning, both dived suddenly and simultaneously; they
emerged about forty yards apart, swam rapidly in the forward
swimming-attitude towards each other, and shook about fifteen
times. After a couple more short bouts, one of them (sex ?)
drove off a third bird which had been following at a little
distance, then came back and shook with its mate. Another
short bout with ruff down, and then they rested for some time,
waking up once to give four languid shakes. A solitary hen
suddenly called some way off; my cock roused himself and
advanced towards her. She first retired, but then came towards
him, and they began to shake. This did not last long, however,
for the cock’s rightful mate dived and came up between the
two that were shaking. She drove the stranger away, and then
came back and had a bout of twenty shakes or so with her mate.
After a time of resting they had a longish bout—some twenty-
five shakes—and as before turned back to back. This time the
procedure was again altered. When 20 or 25 yards away, the cock
dived; the hen waited for him, with her ruff down. He appeared
after nearly half a minute, with some weed in his bill, a little
closer to the hen. He approached, but dropped the weed before
getting to her, and they only indulged in a short bout of shaking.
They then rested and swam to and fro, till a solitary hen
appeared near by, when my cock went up to her. They started
to shake, but suddenly the cock changed his attitude and drove
the stranger away. ‘This change of front was probably due to,
and certainly coincident with, the approach of its rightful mate.
Then both swam off in one direction. The cock, who was leading
by about 50 yards, went close up to a solitary hen who was
ealling, but at the last moment turned and swam back to his
mate, with whom he had a short bout of shaking. After a long
spell of swimming about and fishing, they called to each other,
approached, and had a long bout of.shaking, with less habit-
preening than usual. Again they ended by doing the “ back-to-
back” trick, and again they dived simultaneously. This time,
however, when they came up (some twenty yards apart), though
the hen had nothing, the cock had a very large bunch of weed
in his mouth. They swam together and went through the
regular penguin-actions, he shaking the weed from side to side.
Unfortunately, here again it could not be seen what eventually
happened to the weed. When they settled down on to the
water they did not shake, but separated and swam off together
for fifty yards or so. Then he dived; she followed suit after two
or three seconds. They came up about twenty-five yards apart,
he once more with weed, she once more without it. They swam
together, but he dropped the weed when only a couple of feet off,
and all they did was to shake for a short time. They then went
off fishing, and were lost sight of.
COURTSHIP OF THE GREAT CRESTED GREBE, 553
This pair is also interesting in various ways. The “ back-to-
back’”’ position was never seen in any other birds. Here, too, the
pair itself and both the individual birds of the pair had well-
marked idiosynerasies. ‘The very frequent bouts of shaking, the
several ‘ flirtations” of the cock, and the fact that the cock
brought weeds up three times (the hen not at all) are all worthy
of note. The twice-repeated dropping of the weed just before
the cock reached the hen is very curious; perhaps the hen’s
having no weed had something to do with it.
Once more, too, it is seen that all these emotional actions may
take place far from the nest, and so without any direct relation
to the act of pairing.
13. April18. 2.50-3.35 p.m.
Two birds, one certainly a cock, the other doubtful, were
swimming about and fishing, 10 to 30 yards apart. Each
frequently went into the Dundreary attitude and barked,
apparently at the other, the obvious cock less frequently. Once
the doubtful bird dived and came up a dozen yards or so from the
other; both advanced a couple of yards, stopped, regarded each
other for some moments (rather fiercely, it seemed to me), and
then retreated. I watched them for three-quarters of an hour,
and their general behaviour was the same throughout, except
that they ‘‘ Dundrearied ” less frequently as time went on.
I do not understand the relation between these two birds. I
think it was a hostile one; possibly they were two cocks on the
borders of their respective territories and jealous of their
_ frontier rights.
14. April19. 6.30-6.40 p.m. Close to the reeds.
Here was a curious little “‘ domestic drama” :—
A. pair was swimming about together, and a solitary hen with
a very small ruff was not far off. She was obviously very much
wanting some emotional excitement, for she kept on swimming
up towards the cock, especially when the other and ‘“ rightful”
hen was some way off. ‘he lone bird would swim up to within
three or four yards, eagerly, yet nervously, then turn and go off
as if frightened. The cock was rather indifferent ; once or twice
he began swimming after her, but never got far. Once he was
left mid-way between the two hens, and behaved exactly like the
legendary Ass between the two equidistant bundles of hay. He
looked first one way, then the other, back and forth, back and
forth. At length his mate came a bit nearer, and he at once
turned and swam towards her.
At length the cock’s mate dived: the ‘“‘ wrong” hen at once
dived too, and when she came up found the other hen between
herself and the cock. After a minute or so she approached again,
and this time all three birds dived several times, and finally the
single bird was driven right away.
5d4 MR. J. S. HUXLEY ON THE
Throughout, curiously enough, there was no sound, nor any
erection of crests, nor any going into Dundreary-position ; all
three birds stayed always in the normal swimming attitude.
Whether the late hour had anything to do with this I cannot say.
The diving and driving away, however, showed that there was
some very real jealousy aroused.
It is worth noting that the second time his mate dived to drive
the strange hen away, the cock joined her, but not the first time.
15. April 20. a.m.
I noticed two birds shaking vigorously. At the close of the
bout, a third bird—a hen—came slowly up to the shakers from
where she had been resting some twenty yards away. As she
came nearer, the cock seemed to look with some hostility at the
bird with whom he had just been shaking. This bird, as the
event conclusively proved, was only a stranger, and the hen that
had been approaching was his rightful mate. The ‘‘right ” hen
then swam at the “ wrong” one and drove her away (no flying or
diving); then she turned and swam towards the cock. When
still three or four feet apart they started shaking, in the excited
forward shaking-attitude, with ruffs well up. There were about
twenty shakes; as the bout went on the birds lapsed into the
ordinary shaking-attitude, The strange hen stayed close by, but
after the shaking was over, the cock’s rightful mate swam at her,
and there was “‘ confused diving,” eventually ‘ involving ” all three
birds, aud ending in the odd bird being driven off.
The stimulus given to emotional excitement by jealousy is here
well brought out; but it is curious that the cock’s true mate,
although so close, did not interfere until his bout of shaking with
the stranger was over and done with.
11. MisceLLAngEous Novres.
1. Fishing.
The birds may often be seen to pick small objects off the surface
of the water, often going on for a considerable time. These
objects seem certainly to be eaten, but what they are I do not
know. This habit does not appear to have been previously
noted.
One bird which I saw fishing by itself for over an hour (a
very long time for a Grebe to do anything continuously) had a
curious habit of putting its head right down into the water
with bill vertical, till the eyes were just covered. It once
stayed like this for a good quarter of a minute, but usually it
took its head out after a few seconds, then after a few more
seconds put it in again, and so on, all the while swimming slowly
forwards. It was apparently looking for prey. I saw it dive
while its head was below the surface, but it once dived from the
normal position—7. e. without any preliminary searching. In
other Grebes I have never noticed this habit.
When fishing, they often go along the bank and look for shoals
COURTSHIP OF TIE GREAT CRESTED GREBE. 55D
of small fish, for when they dive for food close in shore one often
sees swarms of little silvery fish spring into the air all round.
2. Relations with birds of other species.
When a Grebe is on the nest, it resents the too close proximity
of other birds. One hen that I saw sitting was twice annoyed by
Moorhens coming too close; she raised herself from the resting
position, bent her neck forward, and definitely (though rather
shghtly—about to the ‘‘ half pear- shaped position”) erected her
ruff (I am doubtful as to her “ ears”). Once she was silent, but
once she gave a low rasping note. On this the Moorhens
retreated.
This same hen was also roused from her snoozing by the call
of a Coot near by; their sleep on the nest must be very light.
Onee, on the open water, I saw a hen Tufted Duck happen to
come close by a Grebe. The Grebe, strange to say, seemed
greatly alarmed, flapped off for some yards, and dived, regaining
ealm again when the surface was regained at a safe distance.
When neither bird is sitting, the Grebes’ nests and platforms are
often ascended by other birds. Several times I have seen Moor-
hens climb on to nests with covered eggs, peck about for food,
and swim off. On what was probably a pairing-platform I once
or twice saw a pair of Wild Duck, and several times a pair of
Shovellers ; they were enjoying a comfortable nap !
On one occasion a Maliard was seen on a nest: a Grebe came
along with a mouthful of weed for the nest, and at its approach the
intruder hastily got off; the Grebe, bower. , pursued him for three
or four yards, before turning and laying the weed on the nest.
3. The Grebe essentially diurnal.
In the spring of 1911 I had been on the Welsh coast, where
> there was an abundance of shore-birds. Of these, certainly the
Redshank, Sheldrake, and Curlew (and very likely others) seemed
to be as wakeful by night as by day, and the special and unmis-
takable courtship-notes of the first two species were heard all
night long, especially on moonlight nights.
My brother and I each slept out a night at the reservoirs to see
whether the Grebes behaved in a similar way. During the dark
hours, however, there was nothing to be seen or heard of the
ines. In the early morning, at the first faint showing of the
false dawn, a few Grebes began to call, and various other birds,
too, showed signs of activity. Unfortunately, between this and
actual sunrise I fell asleep again. My brother, however, watched
the whole period without noting anything of interest. Pyeraft
(11) finds the same hold good in September. Selous (’01) says
that, like many other birds, the Grebe is most active in the first
hour or two after it has become light.
The Crested Grebe is thus a pur ely diurnal species : such birds
as Owls and Nightjars are purely nocturnal (or perhaps late-
Proc. Zoou. Soc.—1914, No. XX XIX, 39
556 MR. J. S. HUXLEY ON THE
crepuscular would be more accurate, especially for the Nightjar,
who is silent for at least the three or four midnight hours);
while in some species, like the Redshank (or the cock Nightingale
in spring), periods of activity and rest alternate throughout the
twenty-four hours.
4. Rest and reserve.
In spite of its being active only during the daylight hours, the
Grebe spends many of these resting, in the attitude so well
described by Selous as resembling a pork-pie (see Pl. I. fig. 2).
This is but one further instance of the principle of reserve that
runs through all life. In watching birds two forms of this are
especially brought under notice—the reserve of time and the
reserve of nervous energy that are present in normally favour-
able conditions.
In the Grebe, the many hours of the day spent in sleeping,
or at least in what Sidis calls the hypnoidal state, represent
the time-reserve. These extra hours of sleep, of course, in-
crease the energy-reserve. This latter is,in most birds, got rid of
in actions which seem entirely without biological significance—
they merely excite pleasure by releasing the energy in bodily
movement or in sound: think of the pleasure-flights of gulls in
early spring, or of swallow-broods in late summer (here accom-
panied by twitterings), or the antics of wagtails in fine autumns
on the lawns. In the spring, however, the surplus energy of
many birds has been seized upon by Sexual Selection, and used
up in fighting or in display (ef. Wallace's general ideas on the
role of energy in Sexual Selection, and Howard (713)).)
the Grebe, similarly, it has been diverted into fresh channels
through Mutual Selection, and thus pressed more directly into
the service of the species.
5. Powers of learning by experience.
As an interesting side-light on the psychology of these birds, I
will record an incident seen by my brother.
A Grebe had caughta very large fish and was trying to swallow
it in the usual way, first throwing its head violently back and
then stopping it suddenly, thus jerking the fish (which, of course,
is held so that its head was foremost) down the throat. This fish
however, was too big. After a long period of fruitless jerking,
the bird was forced to put the fish out into the water. The fish,
being still alive, swam off. This was too much for the Grebe,
who at once dived, caught it again, and again attempted i
swallow it. Naturally it had again to put it out and the whole
process was repeated. It attempted to swallow the fish four times :
the fifth time it let it swim off as best it could. The whole thing
is thus a reflex chain: ‘“ See fish—catch fish—try to swallows
use—put fish out:—See fish....” and soon. It did, however
profit by experience, for each time it made rather fewer efforts th
swallow it, and at last stopped its fruitless trying altogether,
COURTSHIP OF THE GREAT CRESTED GREBE, NG
6. Calls.
I give a brief list of the chief calls employed by the Grebe. As
we should expect in a bird with such a complex emotional life,
different calls are used in different cireumstances—we have in
them another method of expressing emotions.
(i.) Zhe groan.—Typically, a deepish, fairly loud groaning
sound, not guttural or rolling ; occasionally it was given on a
somewhat higher note, and then “ rolled ” slightly.
This is not acommon call: I only heard it coming from birds
in the reeds. I do not know with what emotions or actions it is
associated.
(i1.) The bark.—This is given when one bird is searching for its
mate (first stage of Discovery ceremony), or as an indication of
hostility towards a bird of another pair; and, I think, on no other
occasions : it seems, also, only to be uttered when the bird is in
the Dundreary attitude. There are two chief variations -—_
(2) A loud, rolling, rather shrill and “trumpeting ” bark,
several times repeated.
() A much less loud bark, not so long, not shrill at all, nor
rolling ; repeated quite quickly, but only a few times.
(1i1.) Zhe shaking call.—This is only given during a bout of
shaking. I believe the two are always associated (except
perhaps at the end of very long bouts). It is especially marked
when shaking takes place in the “ forward ” position.
It consists in a rapid alternation of two sounds on two notes —
a consonantal sound—4’p or #c—on the low note, and an indefinite
vowel sound about a tone higher.
(iv.) Zhe “ Double Trumpet.”—This is a very strange-sounding
call, generally given from the reeds, often when near the nest.
It is somewhat of the same timbre as the love-call of the Tufted
Duck, but lower, louder, and more throaty—being, indeed,
extremely guttural.
It is composed of two halves, with a slight pause between.
The first half is something like ah or aw, three or four times
repeated, and rises very slightly. The second half may be repre-
sented thus :—swaa-aa-ah: it sinks rapidly, and is as it were
pressed out, being loudest in the middle.
(v.) The “ Dentist-cull.”—I have given it this name as it re-
minded me irresistibly of that rotating instrument of torture
used by dentists for boring. Imagine the biggest burr grinding
very stiffly for a few seconds, then suddenly running more freely
and whirring for a little. This will give a very good idea of the
sound: and this double sound is repeated several times. I have
only heard it from the reeds: it often follows the “« groan.”
(vi.) Zhe Owl call.—This very much resembles the common
“ker-wick” of the Brown Owl, except that the initial k and w are
not given by the Grebe. It may be repeated, and is often heard.
Perhaps it is the simplest, least definitely emotional call-note
(recognition-note).
39*
558 MR. J. S. HUXLEY ON THE
7. Territories.
For a general review of the subject, see H. Eliot Howard (13).
Like most (or all) monogamous birds, each pair of Grebes
appears to stake out for itself a definite region or territory,
from which intruders of the same species are jealously driven off.
The Grebe, however, differs from birds like the Warblers, and
from the Kingfisher. Such birds live almost exclusively in their
own territory during the whole of the breeding-season, feeding,
sleeping, courting, and nesting in it. With the Grebe, on the
other hand, the territory (to judge from my own experience and
from certain of Selous’s observations) is a comparatively small
piece of water in the vicinity of the nest, and therefore near the
reeds. The open water and the shore, when bare of reeds, is
‘Common Land,” so that almost all the fishing is free to all.
And thus, as a matter of observation, nearly all the feeding and
nearly all the courtship of the birds takes place on this common
ground. This fits in very well with the fact that all the hostilities
I have ever seen on open water were apparently always due to
sex-jealousy. It is only in respect of nesting, of pairing and
the pairing-ceremonies (and probably of sleeping) that the birds
restrict themselves to their territories.
So far as the relation of food and territories go, one might
draw parallels between birds and man; the affairs of the Warblers
would correspond to (present-day) agricultural conditions, the
Kingfisher gives us riparian ownership of fisheries, while the free
deep-sea fisheries are represented by the common open-water
of the Grebe.
8. Swimming Abilities.
Two feats of skill call for notice. In the first place, I have
twice seen birds swimming forwards, in a comparatively straight
line, and apparently with intention, while their heads were tucked
away under their wings.
In the second place, I have seen a Grebe, when frightened off
her nest, dive and swim a good forty yards under water before
rising, although the water was so shallow that she made a ripple
on the surface all the time, and so overgrown with reeds that the
bird’s course had to swerve continually round the obstacles.
9. Stretching of Wings.
Every bird-watcher must be familiar with the habit of Cor-
morants and Shags, of holding their wings out from the body,
apparently for the purpose of sunning them. I have observed
this in the Grebe, but curiously enough only in one bird, which
acted thus twice in ten minutes. It had been preening itself,
and suddenly, raising its anterior end slightly, it stretched its
wings horizontally. They were much arched, and showed the
COURTSHIP OF THE GREAT CRESTED GREBE. 559
white bars very distinctly. After 15 or 20 seconds, during which
I think a little further preening was done, the wings were brought
back to the normal position.
12. Postscript.
Owing to accidental circumstances, it was unfortunately only
after the completion of the MS. of this paper that I was able to
read Mr. W. P. Pyeraft’s interesting book on the Courtship of
Animals (Pyeraft, 713).
As Lintend to attack the problem of the relations of Mutual
and Sexual Selection in a more general article, it will be un-
necessary to discuss bis general conclusions here m detail. Let
me only say that had I, before writing this paper, read his general
discussion of female choice and of the modifications required in
Darwin’s original Sexual Selection theory, much of my own
theoretical conclusions would have been differently expressed,
although perhaps not essentially altered.
Let it be particularly noted, however, that Mr. Pycraft himself
is careful to point out that Darwin’s main conclusions stand firm.
As I understand it, the chief modification necessary relates to
female choice. Display and ornament do not act on the esthetic
sense of the female, but on her emotional state; they are—using
the words in no narrow or unpleasant sense—excitants, aphro-
_ disiaes, serving to raise the female into that state of exaltation
and emotion when alone she will be ready to pair. This is
brought out most vividly in the nuptial behaviour of the Newts
(Pycraft, “13, p. 170). No one, after reading this, can fail to
understand not only that the pure Darwinian theory needs
modifying, but also the direction in which it must be modified.
But the element of choice does, in another form, remain. In
animals such as Birds, where there is a regular pairing-up season,
and where, too, the mental processes are already of considerable
complexity, it is impossible to doubt but that mating may be, and
in some species is, guided by impulse, unanalysable fancies, indi-
vidual predilection. There, in a rudimentary state, we find that
form of “ choice ”—intuitive, unreasoned, but none the less im-
pervious, and none the less in its results a true choice—which
reaches its highest stage of development in the intensely-felt
affinities of man and woman—in that condition known as “ falling
in love,” where the whole of the subconscious mental activities
become grafted on to the inherited sexual passions, the whole
past of the mental organism is summed up in the present, in
the intensely real act of choice which chooses one from among
thousands and says, whether in words or no, “that one being,
and no other, is the being that I desire for my mate.”
That a choice of this type can exist in birds is shown by the
subject of this memoir. ‘The individual variations in the court-
ship-actions provide the raw material for preferential mating,
and the fact that the birds of a pair often both show some
560 MR, J. 8S. HUXLEY ON THE
special variation in the form of action, in itself proves that
such preferential mating may and does occur.
The first modification of Darwin’s ideas leads to a second—to a
modification of the way in which Sexual Selection works. If
display is normally an excitant, then there is no ‘“ need” for a
preponderance of males, nor for actual rivalry between several
males. Selection is primarily a matter of level. The female does
not choose the “ best” out of a bunch of suitors ; but those males
in which the ornaments and display-habits do not reach a certain
standard, will not be able to raise the female’s emotional state to
the requisite pitch, and so will die without offspring. (‘This state-
ment as it stands goes too far; it will serve, however, for the
time, to show the general idea.)
This primitive condition has been modified in two ways (and
on the existence of these two quite distinct lines of development
I feel Mr. Pycraft has not laid sufficient stress). In the first
place, in polygamous and polyandrous (and perhaps “ promis-
cuous ”?) species there may bea rivalry between several males
in the presence of the females, as in the Ruffand Prairie Hen.
Jn the second place, the line of mutual selection was started.
Whether in origin mutual displays too acted as excitants, it is
not yet possible to say until more observations are at hand. That
such may be the case, is possible from some observations of my
own on the Herring-Gull. At any rate, in specialized forms of
this form of courtship, such as that of our Grebe, this excitatory
function is completely in abeyance.
With Mr. Pycraft’s insistence, first, on the principle of Ortho-
genesis and its importance for the origin of sexual (and other)
forms of ornamentation, and secondly, on the necessity for a
psychological pomt of view in our interpretation of the courtship-
phenomena of animals, I am in entire agreement.
List or LrreraturReE.
713. Howarp, H. Exior.—The British Warblers. Parts 1-8.
Londen, 1907-13.
*12,, Huxuey, J. S.—A First Account of the Courtship of the
Redshank (Yotanus calidris Linn.).
P. ZS. 1912, pp. 647-655.
Z12.. " 55 The Great Crested Grebe and the idea
of Secondary Sexual Characters.
Science, n. s. vol. xxxvi. pp. 601—
602.
"11. Pycrarr, W. P.—Habits of the Great Crested Grebe.
Bield, vol. exviii.- Oct.. 7, @l9ii
pp. 823-824.
ES, ° The Courtship of Animals. Hutchin-
son & Co. London, 1913.
COURTSHIP OF THE GREAT CRESTED GREBE, 561
701. Senous, E.—An Observational Diary of the Habits—mostly
domestic —of the Great Crested
Grebe (Podicipes cristatus). Zoolo-
gist, May 1901, pp. 161-183, Sept.
1901, pp.339-350, Dec. 1901, pp. 454—
462, Apr. 1902, pp. 133-144.
02. i (Reversed pairing in Moorhens.) Zoologist,
May 1902, pp. 196, 197.
"13. Pe (Display of Swans.) Zoologist, Aug. 1913,
pp- 294-313.
Screntiric NAMES OF BIRDS MENTIONED IN THE TEXT.
To save constant reference to birds by both English and scien-
tific names, and to help foreign readers, T append this list.
Blue Tit. Parus ceruleus L.
Bustard. Otis tarda L.
Coot. Fulica atra L.
Dabchick. Podiceps fluviatilis (Tunst.).
Egret. Ardea, Egretta, Herodias, Garzetta.
Fulmar Petrel. Fulmarus glacialis (1L.).
Guillemot. Uria troile (L.).
Heron. Ardea.
Herring-Gull. Larus argentatus Gmel.
Kagu. Rhinochetus jubatus.
Mallard. Anas boschas L., 3.
Moorhen. Gallinula chloropus (l.).
Peacock. Pawo cristatus L.
Prairie Hen. Tympanuchus americanus.
Razorobill. Alcea torda L.
Redshank. Totanus calidris (L.).
Shoveller. Anas clypeata (L.).
Snipe. Gallinago celestis (Frenz.).
Swan (Whooper). Cygnus musicus Bechst.
Tufted Duck. Fuligula cristata (Leach).
Warblers. Sylviidee.
Wild Duck. Anas boschas L.
EXPLANATION OF THE PLATES.
[The figures were drawn from my notes and rough sketches by Miss Woodward,
to whom I am much indebted for the interest and_ care she has shown. Taken as a
whole, they give a far more graphic and accurate idea of the birds’ general appearance
and behaviour than any other illustrations of which I know. |
All figures refer to Podiceps cristatus.
PruateE I.
Fig. 1. Head and neck, showing ruff and ears relaxed.
2. Resting attitude. Note the position of the head, and the curve of breast
and rump. In most figures these are erroneously represented.
3. Search (Dundreary) attitude. Note the ears relaxed, the crest spread
longitudinally (sometimes it may touch the water).
ON THE COURTSHIP OF THE GREAT CRESTED GREBE,
. Head and neck in Shaking-attitude (ears erected vertically, ruff pear-shaped).
. Shaking-attitude from behind. Note the curious shape of the lower part of
the neck.
. A pair in the Forward (exeited) Shaking-attitude. Note the head bent
down, the neck strained forward; the slope of the body and cock of the
tail are also very characteristic.
. The Cat-attitude (Display). The general attitude is very well represented.
More white should show on the breast; and the dark portion of the
wings should be grey. ‘To represent them black lessens the effect of the
real black on the crest, which in actual life is the central aud most con-
spicuous part of the picture.
. The Passive Pairing-attitude. Note the strange stiff appearance, the humped
b=) i=) )
back, and the total closure of the crest.
Prare II.
. The “ Ghostly Penguin” (attitude on emergence of the diving bird in the
Display Ceremony). Note the head bent down, and the forward curve
ot the top of the neck.
. The same as fig. 9, side-view. Owing to the short time occupied in the
action, Icamnot myself be sure that all the details in figs. 9 and 10 are
accurate. The general appearance, however, is well given.
. A pair shaking. Note the erect necks, and the tails slightly cocked up.
. Display Ceremony : the diving bird just fully emerged. (This is the only
tigure which is not satisfactory. It gives the positions etc. well, but does
not recall reality in the vivid way done by the others.)
. The Penguin Dance. Here again the whole ceremony takes such a short
time that I cannot vouch for details; but the general appearance is very
well suggested.
P Z. S. 1914, PERKINS. Pl. I.
= > =
Ue aay
|
=
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7
lo |4 8
Cn i
l!
15 17
AS aN hat By
\2 16
13
20
22
R.C.L. Perkins, del. Bale & Deamelsacn lc imp -
STRUCTURAL CHARACTERS OF PARALASTOR, SAUSS.
ON THE WASP-GENUS PARALASTOR. 563
34. On the Species of Alastor (Paralastor) Sauss. and some
other Hymenoptera of the Fanily Humenidee. By R.
C. L. Perzins, M.A., D.i8e., F.Z.8.
[Received April 9, 1914; Read May 19, 1914.]
(Plate 1.*)
INDEX.
Ethology : Pages
Paralastor, mimetic colour-groups of ............... 6... 563-565
Geographical :
Paratastor, am Australian Genus) 22. 5.).21..6 02. -20---e---e 563
Systematic :'
Paralastor, division of Alastor Sauss., raised to generie¢
PUN ra Seas raeea sore Gedennd ecieromae hacen Se 563
A NE WeSPEGCIES Oli seh ecu eoess seen Seek eee) (B79=621
Pseudozethus australensis, gen. et sp. N..........-....... 622-623
JETT OPOCUIO COUSUREHKAD OR S\05 Wo ga3d06 bab 5d00ce 264 05a 464 oo dcr 623
Abispa meade-waldoensis, Sp. 0. .....-...-seveeeee eee ee ees 623
The name Paralastor was given by Saussure to one of the several
divisious of the genus Alastor which were characterized by him.
It includes all the species of the Australian region, except one,
Paralastoroides clotho Lep., and this differs but little from the
other Australian species.
Paralastor is a valid genus, and has no close affinity with the
non-Australasian forms included in Alastor. Its most remark-
able character lies in the structure of the antennz of the male, the
peculiarities of which were quite overlooked by Saussure, as they
were also in other genera of Eumenide, such as Abispa and
Pachodynerus. The antenne of Paralastor g consist always of
eight well-developed joints, followed by four, three, or two small,
or often very minute, ones. This deficiency in the normal
number of joints of the male is always accompanied in both sexes
by a thoracic structure, so different from that of the Alastor of
other countries that the genus is extremely well defined. The
thoracic characters have been so clearly stated by Saussure that
it is not necessary to repeat them here.
The 8. African species Alustor brawnsi Meade- Waldo, and Alastor
variolosus Bingham from Ceylon are members of one group,
quite distinct from Paralastor, having ordinary male antenne
with the normal number of joints (13), as in Odynerws and like
European Alastor. On the other hand, the E. African Khynch-
alastor fuscipennis M.-W. is more related to the S. American
species, such as A. melanosoma Sauss., but the clypeus is pointed
at the apex and the 2nd ventral segment of the abdomen has a
costate sulcature followed by a deep large impression, and is not
merely flattened between lateral carinations.
The coloration of the species of Paralastor is interesting, in
that, like the Odynerus of the Hawaiian Islands and elsewhere,
* For explanation of the Plate see p. 624.
564 DP. R. C. L. PERKINS ON THE
the species form a number of distinctive colour-groups, which are
quite different from the groups formed by them when characters
of structure are considered, Particularly interesting are the
species of the second division of the genus as arranged by me,
since several of these closely resemble small species of Australian
Icaria, social wasps, which are found in the same localities.
‘he resemblance is rendered far more perfect from the fact that
in these Paralastor the basal abdominal segment is reduced in
size and is less wide than usual, so that the abdomen approaches
in form that of some of these social wasps.
As to the other division of Paralastor, the species not only form
colour-groups amongst themselves, but these groups mostly, and
perhaps all of them, reproduce the colours and patterns of other
genera of solitary wasps, Odynerus, Rhynchiuvm, and Abispa.
Even the bees of the genus Hylaeoides (Prosopide) are drawn
into these colour-groups, and both in these bees and some of the
Odynerus the resemblance extends beyond the matter of colour,
so that certain peculiarities of structure are reproduced in all. I
have elsewhere (Ann. & Mag. N.H. (8) ix. 1912, p. 108, Hylacoides,
and p. 121, Abispa) referred to cases of this structural mimicry
combined with mimicry in colour. How great is the resemblance
may be judged from Saussure’s description of Alastor fraternus,
of which he says: ‘‘ Presque identique par la coloration avec les
Odynerus clypeatus et concolor; formes les mémes,” ete.; but this
case, naturally, does not compare in interest with that of the bee
HHylaeoides, where structures identical with those of Paralastor
are assumed, these structures being unique in that family of bees
to which Hylaeoides belongs. In the case of the Odynerus and
other Eumenid genera, where structural and colour mimicry are
combined, we are at least dealing with members of a single
family, however distinct the genera themselves may be.
All the species of Paralastor from Tasmania that I have myself
seen, are extremely similar in appearance, with nearly white
markings and two very narrow abdominal bands. Odynerus of
quite similar appearance were taken with these by Mr. R. E.
Turner last year.
In N. Queensland, P. optabilis, which has a coloration entirely
unlike any other Queensland species of its genus, is mark for
mark the same as an Odynerus occurring with it, and both have
similar structural peculiarities.
At Adelaide in 8. Australia and in Victoria are numerous
species with deep dull orange or red markings, superficially either
entirely or nearly resembling one another, though falling into
the most diverse structural groups. These species seem to vary
in their patterns individually, and it would appear that different
species exhibit similar variations of pattern.
Orange-marked species (though found elsewhere) are abundant
in Queensland, while W. Australia has a very distinct colour-
group, the thorax of the species being largely red, and the basal
abdominal segment similarly coloured, while the rest is black. It
WASP-GENUS PARALASTOR. 565
is interesting to note that the four members of this last colour-
group that are at present known fall each in a totally different
group structurally. Of course, W. Australia has species other-
wise coloured, which resemble those from other parts of the
country.
North Queensland has a number of species remarkable for the
ornamentation being nearly confined to the large 2nd abdominal
segment, the pattern of this being very distinctive, sometimes
ferruginous, sometimes yellow or nearly white.
It is evident that many and perhaps most of the species exhibit
considerable variety in colour or pattern of colour, as also in
smaller details of structure, but to what extent this is the ease,
whether as regards individuals from one station or those from
separate localities, the material at my disposal is far too limited
to decide. This material consists of the British Museum Collec-
tion, excluding the actual types of Saussure and Smith (most of
which, however, I have examined), of the collection in the Hope
Department of Zoology in the Oxford Museum, including types,
and of my own collection. The total number of specimens is
only 333, although 2000 or more would hardly be adequate for
dealing with the genus satisfactorily. Moreover, as six species
together account for 101 of the 333 examples, it will be seen that
about 90 species are represented very poorly indeed. There is
no doubt that great numbers of species and local forms remain
to be discovered, and the genus will ultimately prove to be very
difficult. Even at present the species are not at all easy, com-
pared with the Hawaiian Odynerus, which about equal them in
number.
A great deal of the material in the British Museum and at
Oxford is very old and often dirty, and has, further, been badly
mishandled in the desire to extend the wings.
In Alastor, Odynerus, and kindred genera, the abdomen should
always be kept flat on the ventral surface, and not bent at the
suture between the first and second abdominal segments, as is often
the case. It should, however, be bent downwards at the petiolar
articulation, so as to fully expose the propodeum. If properly
mounted in this way, the base of the second ventral segment will
be fully exposed and the structure of the segment readily observ-
able. Tio anyone who has handled many hundreds of species of
Kumenidz, proper preparation of the specimens is of minor con-
sideration, as he will easily make allowance for the retraction or
flexure of the second ventral segment, but to one not so circum-
stanced, the difference in appearance caused by poor preparation
may be a source of serious error.
Two species which I have described on single specimens have
not the neuration of Paralastor, but of Odynerus, the second
cubital cell not being petiolate. Whether they are constant in
this character or not, remains to be proved. It isin any casea
very feeble one, for in some species the petiole of the cell is very
much reduced, so that it was quite certain that sometimes, either
566 DR. R. C. L. PERKINS ON THE
as a variation or as a specific character, it would be altogether
wanting. The importance placed on small differences in neura-
tion in Hymenoptera has been much overrated, and there is little
to commend these characters, excepting the fact that they are
obvious at a glance.
Some described species it is impossible for me to tabulate, as I
have never seen specimens of them, and the descriptions, admir-
able as Saussure’s are in general, omit certain important points.
Some, not included in the tables, I have referred to subsequently
with regard to their probable position. .The following may be
brietly mentioned here :—
P. albifrons Fabr. If the tegule have a coarse, deep, and con-
spicuous puneturation, this species would appear to be
extremely closely allied to P. habilis and subhabilis, but
superficially distinct by having only ‘“‘ deux trés petits points
orangés sur le prothorax” and the apical band of the 2nd
abdominal segment semicireularly emarginate, the emargina-
tion being much deeper in the other two.
P. australis Sauss. This would fall in my table in the group of
P. princeps, oloris, ete. If Saussure’s specimen was correctly
stated to be a female, the yellow clypeus and front of the scape
of the antennz would distinguish it from the females known
to me, which resemble it otherwise in colour, as well as the
spot on the mesopleura from either males or females of this
section.
P. clotho Lep. I know no species with this pattern, except
Alastor unifasciatus Sm., which lacks the scutellar spots, has
dark tegule and very dark wings with violet iridescence,
and is not an Australian species.
P. cruentatus Sauss. I think it is impossible to identify this
species without seeing the type.
P. flaviceps Sauss. I have included this in my table where it
would appear to belong. If the tegule are coarsely pune-
tured it is correctly placed. In any case it is distinct
superficially from all the species which have an entirely
black basal segment by the sulphur-yellow head with a black
oval mark on the vertex, enclosing the ocelli.
P. graefei Sauss. This species from Ovalau is unknown to me
and not included in my table.
P. lachesis Sauss. Not included in my table, as the sculpture of
the tegule is not indicated in the description. Superficially
it resembles P. orientalis P.
P. lateritius Sauss. I suspect this of being a slight colour-variety
of P. carinatus Sm.
P. nautarum Sauss. Probably tabulated rightly; as it is one
of the species with “un tubercle saillant en dessous du
deuxiéme segment,” an emarginate clypeus, and the abdo-
men all black, except the orange basal segment, it cannot be
confused with any other known to me.
WASP-GENUS PARALASTOR. 567
P. smithi Sauss. I unfortunately failed to examine the type of
this species, which is in the British Museum, and I cannot
place it in my tables from the-description nor identify it
with any of those described by me.
P. hirtiventris Cameron. IhaveseenCameron’s ¢ type. Itisthe
largest species of the genus, the tibie with long conspicuous
hairs beneath. The second ventral segment is greatly raised
behind the suleature, prominent at the top of the truncation
medially, and clothed with long erect hairs all over, the
following segments densely hairy. The thorax is all black,
the 1st and 2nd abdominal segments with wide orange apical
bands clothed very conspicuously with erect hairs.
T have to thank the authorities of the British Museum for the
loan of their specimens for description. Several of these bore
type-labels of Saussure’s, but the species were not deseribed by
him. I have, except in one instance, adopted the names applied
by him to these species. Prof. E. B. Poulton very kindly sent
me the whole of the Hope Museum collection of this genus.
To Mr. Meade-Waldo I am particularly indebted for examining
special structures in some of the British Museum type-specimens,
which I had overlooked in the limited time at my disposal there,
and for sending me descriptions of some others which I needed
to complete this paper.
In the following descriptions the measurements are always
taken from the front of the head to the apical margin of the
2nd abdominal segment.
In my description of the 2nd ventral segment I use the adjec-
tive “elevatus” as applied to the part which lies behind the
transverse sulcus, because it is convenient and natural to reverse
the insect in examining this segment. Of course in reality the
segment is not raised, but produced downwards behind the
sulcus.
When a species is contained in more than one of the three
collections examined by me, the actual type-specimen has always
been selected from the British Museum Collection.
The genus Paralastor may be divided into two main sections,
one of which contains by far the greater number of species and
exhibits by far the greater variety of structure. The species of
the small division (the Divisio secunda of the following table)
may be characterized as follows :—
Species semper graciliformes ; segmentum primum abdominale
conspicue parvum, nunquam fortissime transversum, sed sub-
campanuliforme, secundo semper, basim versus, utrinque con-
spicue angustato. Segmentum 2 dorsale basim versus fortissime
convexum, ita ut semper (a latere visum) supra segmentum
basale fortiter aut fortissime elevatum appareat; secundum
ventrale, insecto resupinato, post suleum suum transversum
semper supra partem suam basalem (sive presulcalem) fortissime
568 DR. R. C. L. PERKINS ON THE
oblique elevatum, elevatione summa media nunquam antice pro-
ducta aut tuberculiformi. Propodeum (excepto P. picteti, cujus
propodeum nigrum est, abdominis segmento primo rufo, postice
pallide-marginato) colore signatum.
There is no absolute discontinuity between the two sections,
and one or two species I have placed in both tables. For the
most part, however, the insects of either group are separable at
a glance by the characters above given, and since the one or
two dubious forms are included in both tables, I do not think
there can be the least difficulty in placing any of the known
forms.
It is clearly the small size of the basal segment that gives the
species in the second division their characteristic superficial
appearance, and by this also their resemblance to some of the
small Australian species of /caria is made much more perfect.
Ali species that have not the above characters are necessarily
referable to the Divisio prima.
Divisio prima.
Thorax, pronoto excepto, semper totus niger ; abdominis seg-
mentum basale aut totum nigrum, aut linea aut fascia
subabbreviata, hand ad angulos ipsos laterales extensa, sed
ante hos evanescente, decoratum.
(Clypeus semper emarginatus, et abdominis segmentum
2 dorsale basim yersus haud conspicue convexim
elevabum))...2%os-aeae wh atesiaecms pocaces toe ea eda oe eee eee 1
Thorax spe aliter decoratus ; segmentum abdominis basale
nunquam totum nigrum, sed fascia apicali integra, aut non-
nullis in speciebus ex majore parte colore ornatum ............ 16
1. Ale anteriores tot, aut fere totw, nigricantes aut profunde
infumatz et conspicue ubique cruleo-nitentes; clypeus
utrinque carina longa, bene exstante, ad aut post medium
extensa, haud munitus.
(Abdominis segmentum primum totum nigrum) ............ 2
Alz anteriores in A. wnifasciato Sm. solo ubique violaceo-
nitentes, quo in casu carinis longis, post medium extensis,
Glypeusestprseditus, 136: 2ie tac as on une eit ee eee oe ee 4
2. Abdominis segmentum 2 rufo-brunneum aut brunneo-auranti-
acum, triangulo elongato, nigro, mediano, a basi post
medium segmentum extenso, ornatum ............... saussuret, Sp. Nn.
Abdominis segmentum 2 nigrum, fascia apicali flava, rufescente,
aut aurantiaca, haud basim suam utrinque attingente,
GUI HULA ee fe gae toe aca ONCE er eters MeCN CP OE he 3
3. Femora, trochanteres et tibie nigra; fascia segmenti 2 ab- ~
dominalis antice emargimata ..................0..00.00s infernalis Sauss.
Femora, trochanteres et tibie rufescentia; fascia segmenti
2 abdominalis angusta, perpaullo utrinque latior... rufipes, sp. n.
4, Clypeus fortissime emarginatus, emarginatione, ac semicirculus,
zeque profunda, aut etiam hoc profundiore ..................... 5
Clypeus haud profundissime emarginatus, emarginatione arcum,
Semi(cireul OMMINOLeMestOrmanle eee teee eee eee eee aan eee a
5. Ale anteriores et posteriores fere squaliter profunde in-
fuscatze (parte costali nigriore); abdominis segmentum
2 totum rufo-brunneum, aut brunneo-aurantiacum, margine
postico primi angustissime, in parte media, cum secundo
CONCOLOLE Brena in meee Se ene yas See eae Fraternus Sauss.
10.
Il
12.
13.
14.
15.
16.
WASP-GENUS PARALASTOR. 569
Alz haud distincte ubique fortiter infumate, sed ex magna
parte hyaline aut subhyalinz ; abdominis segmentum secun-
dum s#pius nigro aut fusco colore variegatum ; si unicolor
est, margo segmenti primi posterior haud coloratus ......... 6
Abdominis segmentum 2 ad apicem suum sat late nigrum,
parte basali (triangulo mediano parvo excepto) colore albo-
flavo ornata, fasciam, profunde angulariter postice excisam,
POWMAMLC MRR estes eect ee eet meh eee Ce sinc Secs conspicuus, sp. Nn.
Abdominis segmentum 2 totum pallidum, aut macula nigra aut
fusca elongata mediana, nonnunquam marginem basalem,
nonnunquam apicalem attingente, ornatum, picturatione
variabili ; color segmenti, ut apparet, testaceo-flavus, aut
pallide ferrugineus (an post mortem discolor?) ... duwbiosws, sp. n.
Stigma alarum nigrum aut atro-brunneum; tegule minu-
tissime punctate, nonnunquam in disco puncta pauca
grossiora ferentes, aut plus minusve leves, et parce obso-
letim punctate; clypeus aut duabus carinis longis, bene
elevatis, conspicue instructus, aut obscurius tricarinatus ... 8
Stigma alarum pallidum ; tegule grossissime et conspicue, seepe
rugosim, punctate ; clypeus carina utrinque conspicua, bene
elevata, carens (deflexione tamen clypei laterum marginem
acutum nonnunquam formante) nec obscurius tricarinatum . 13
. Alz anteriores et posteriores atro-infuscate ; abdominis seg-
mentum 2 dorsale totum ferrugineum, aut nigrum fascia
apicali flava aut aurantiaca ornatum ........................ 20.205 g)
Ale haud atrz, anteriores sole in parte costali atro-infus-
cate, ceteris partibus hyalinis aut leviter infumatis ;
abdominis segmentum secundum aliter picturatum ............ 10
Alz haud vyiolaceo-colore conspicue nitentes ; segmentum
abdominis 2 dorsale totum ferrugineum aut opace auran-
LOLOL HIG Gena bobh lane oannee Seem oe a NDOG Soe SOR RCE oR Cente Ses cognatus Sm.
Alze conspicue violaceo-nitentes; segmentum abdominis 2
dorsale fascia apicali tantum ornatum ............... unifasciatus Sm.
Clypeus utrinque fortiter longitudinaliter carinatus, carinis
foxutiterrenacupese! evabisn CO) espe beet eee tee erence tee 11
Clypeus obscure longitudinaliter tricarinatus, inter carinam
medianam et laterales depressus, apice leviter emarginato
avixed 1 LOMUDALOR (om) Peneeer ert ceteris orate pincers sence seers 12
Abdominis segmentum secundum fascia basali, latissima,
pallide flava, postice late emarginata, ornatum ... conspiviendus,sp.n.
Abdominis segmentum 2 pallide flavum, plagam magnam sub-
triangularem nigram includens ........................ elegans, sp. Nn.
Segmentum 2 dorsale abdominis subsanguineum, plaga basali
nigra (haud ad segmentum medium extensa) notatum ; seg-
mentum primum spatio lunulato ejusdem coloris apicali
CIBACHEIEAY onan was bb nae nae SoBe eaws on bene SAG Re Ere ens tricarinulatus, sp. n.
Abdominis segmentum 2 dorsale ferrugineum aut aurantiacum,
totum pallidum, triangulo basali nigro et brevi excepto.
tasmaniensis Sauss.
Abdominis segmentum 2 dorsale plaga nigra mediana, a basi
ad medium aut post medium extensa, ornatum ............... 14
Caput sulphureo-flavum, ocellis plaga nigra ovali circumdatis.
flaviceps Sauss.
Caput nigrum, macula utrinque postoculari, et macula inter-
EAT RETAIN, MORIAH o" Aad alee bab beaameuc sueece Sone SEA RR OS SeLeGne eRe MeeS 15
Ad brayesipil es) Tauivaranteehentt 8 ote a dei fan Ae ead on Poe habilis, sp. n.
Tegularum margo exterior pallidus ..................... subhabilis, sp. 1.
Thorax totus niger; abdominis segmenta 2 basalia tota
Hern cinea ene dat nay vale occ Daa Cais Ree Sar apicatus Sm.
SOOGIES milter GOlkorna. 65 Suk ke nhs ete ee ee SES e Nee PARC See sunelee 17
570 DR. R. GC. L. PERKINS ON THE
17. Abdominis segmentum 2 dorsale pallide flavum, aut ochraceum,
macula nigra basali triangulari ornatum, apice trianguli nigri
fere ad medium segmentum extenso, segmentis sequentibus
et margine primi posteriore cum secundo fere concoloribus ;
thorax niger, pronoto solo maculis flavescentibus ornato.
pallidus, sp. n.
Species colore precedenti valde dissimiles .....................-.. 18
18. Pronotum, scutellum fere totum, postscutellum, tegule, cum
propodei lateribus, ochraceo-flava ; abdominis segmentum
primum eotlem fere colore fasciatum, fascia media quadrato-
emarginata ; secundum nigrum, apice vix pallescente (¢).
insularis Sauss.
Species colore preecedenti valde dissimiles ........................ 19
19. Abdominis segmentum 2 ventrale post suleum suum trans-
versum in parte media peroblique aut convexim elevatum,
et ibidem hand aut vix ad altitudinem, quam pars sua basalis,
majorem surgens ; clypeus emarginatus ....................00.: 85
Abdominis segmentum 2 ventrale post suleum suum trans-
versum in parte media sepissime fortissime (aut saltem
distincte) supra partem suam basalem elevatum ; aut si pars
postsulealis partem przesulealem (sive basalem) altitudine
haud superat, a suleco sat abrupte surgit pars posterior, et
clypeus “troncabus estore crass oe nconale ian eaeec cin ae eee 20
20, Abdominis segmentum 2 ventrale, post sulenm suum trans-
versum, tantum ad partis basalis sue altitudinem elevyatum ;
clypets Semper pcm ea iis paeeer a eee te ene 81
Abdominis segmentum 2 ventrale, post suleum suum trans-
versum, sepius fortissime supra partem suam basalem
elevatum, aut semper distincte altius quam pars basalis ... 21
21. Clypeus (lateribus deflexis exceptis) usque ad basim perfecte
deplanatus (nee concayus sive impressus) et parti frontis
Interantennali deplanatze, quam perfectissime, adaptatus ;
postscutellim: simplex, imerme ..........-.......+.)-)s)sesee eee 70
Clypeus basin versus aut ubique plus minusve conyexus aut
rare impressus ; si usque ad basim deplanatus, postscutellum
medium tuberculatum aut spiniferum est ........................ 22
22. Clypeus sat fortiter emarginatus, et nonnullis in aspectibus
paullo inter dentes laterales prominens, ita ut, plus minusve
obscure, quasi tridentatus appareat ...............0.cc0c0eeee eee 63
Clypeuspaliten Porm aus eee esses ree eee none eee 23
23. Clypeus carinis acutis duabus elongatis longitudinalibus, bene
exstantibus sporceci hte. qe.) noe ee 62
Clypeus haud ita carinatus, sed margines deflexi laterales
nonnunquam aciem acutam formant .............................. 24
24. Postscutellum medium spina brevi aut tuberculo distincte
acre Tih 1s peel ae Se a ated HMI ee EES Gl 65
Wostscutellam)/inerme | 7:0..04..4....10..2ccersoe ca 25
25. Tegule perconspicue grosse punctate, marginem versus
exteriorem puncta hee grossa ferentes; clypeus semper
@marcing DUS eberter ast eats) h nt ee acl a ieee ae 52
Tegule sepe minutissime punctate ex parte majore, aut fere
leeves, puncta majora marginem exteriorem versus conspicua
rare ferentes, quo in casu clypeus truncatus est ............... 26
26. Abdominis segmentum 2 dorsale in formam conicam fortissime
et abrupte (oblique tamen) elevatum ; clypeus truncatus ... 79
Abdominis segmentum 2 dorsale nonnunquam fortiter convexim
elevatum, sed haud vere conico-tuberculatum, aut si conicum,
el ypeustesbemancina busses stick te yea 27
27. Abdomen fasciis unicoloribus duabus (una in specie compluribus)
flavis, seepissime albido-flavis, rarius lete flayis, ornatum ;
WASP-GENUS PARALASTOR. 7 Il
fascia prima plerumque lineari, rarissime circiter partem
segmenti dimidiam, desuper aspecta, occupante; fascia
secunda haud majorem quam quintam partem segmenti
secundi occupante; pronoto antice bimaculato, maculis a
tegulis longissime remotis, propodeo nigro ..................... 28
Abdomen aut aurantiaco, aut rufescente colore fasciatum ; aut
fascia una aut altera aut ambabus distinctissime bicoloratis ;
nonnullis in speciebus segmentum primum, desuper aspectum,
totum aut fere totum est coloratum, propodeo nonnunquam
haud toto nigro, maculis pronotalibus nonnunquam ad
tegulas aut fere ad tegulas extensis ..................0...0000005. 34 a
28. Species duabus fasciis abdominalibus ornate ..................... 29
Species unica, fasciis compluribus albido-flavis ornata ......... 33
29. Mesonotum in parte dimidia posteriore aut grosse aut densim
et rugose punctatum; frons inter antennas maculata ;
antenn ey cail-arhiculatcom eee nner ena nee eee nee 30
Mesonotum in parte pesteriore haud grosse punctatum, punctis
inter se remotioribus aut irregulariter dispositis ; frons inter
antennas nigra; antenne ¢ 12-articulate ..................... 34
Aneguli laterales pronoti anteriores distincti, evidenter pro-
minuli; mesonotum opacum, et cum scutello ubique quam
densissime rugoso-punctatum. (Tegule flavo-marginatz.)
: emarginatus Sauss.
Anguli laterales pronoti anteriores parum distincti, haud
promimentes ; mesonoti puncturatio minus densa quam in
precedente, punctis (nonnullis in partibus) imter se
GIS GIT CTI SMe reece teN ene eae ee ere mace cans Ae een Ry 31
31. Tegule, ex magna parte aut tote, testacez, haud flavo-notate ;
abdominis seementum primum postice angustissime lineariter
fasciato, fascia ipsa haud vel sparsissime punctata.
(Abdomen in exemplis recentibus pilis longis ubique
vestitum; dG segmentum ventrale apicale pilis sat
longis erectis vestitum; ad basim suam utrinque
pilis densis erectis preditum, his seepe sub segmento
preecedenteraloditis)hoesete eee oeece eset esete parca Sauss.
Tegule sepe flavo-maculate ; abdominis segmentum primum
postice minus anguste fasciatum, fascia ipsa punctis multis
Conspicuegora ditay eine seas ees tee eee eee creer tien ae eens 32
32. Abdominis segmentum tertium et sequentia (a latere visa)
pilis erectis parce vestita; fascia flava segmenti primi
antice media emarginata;. dG segmentum ventrale ab-
dominis apicale pilis erectis sat longis vestitum; Q
THAN GKOYER INI 2) > = hrs BOA Se BNR At Maa ae aoSiatanoL eee Se seace Pee sp. n.
Abdominis segmentum tertium et sequentia solum tomentosa ;
fascia flava primi haud antice emarginata; ¢ segmentum
ventrale abdominis apicale pilis brevissimis vestitum ;
(OV ST TGOP AINE) ieee deta bedaoSE Dae Gomes saree CBNoRpeT TESMeDROR ee frater, sp. n.
33. Clypeus leviter lateque emarginatus, albo-flavus, macula
mediana nigro-signata ; mesonotum dense distincteque
punctatum, tegulis externe testaceis, postice flayo-notatis ;
alee parum fortiter infuscate (Q) ..................... ordinarius, sp. 1.
34. Abdomen fasciis duabus angustissimis flavo-albis; corpus
totum pilis longis vestitum, forma graciliore; tegule fere
nigre ; abdominis segmentum primum haud fortissime
transversum; clypeus aut truncatus aut vix evidenter
30
emarginatus; antenne J 12-articulate ............ punctulatus Sauss.
34a. Clypeus fortiter aut fortissime, levissime aut leviter emar-
Sinatiis, emMaroinatusbalMenm lates stetdsvooe en svedeescese eee. 35
Clypeus aut recte aut subrotundatim truncatus .................. 73
Proc. Zoo. Soc.—1914, No. XL. 40
ie DR. R. GC. L. PERKINS ON THE
35. Abdominis segmentum 2 dorsale (lateraliter visum) pilis longi-
oribus erectis, sat numerosis, conspicue vestitum ; pronotum
antice binotatum, propodeo nigro, puncturatione mesonotali
confusa, leviter impressa. (Tegule testacee ; anguli pronoti
humerales parum distincti.)
Abdominis segmentum secundum pilis brevibus erectis sparsim
vestitum, aut fere nudum ; pronotum nonnullarum specierum
ad tegulas aut longe has versus coloratum ; propodeo non-
inillempunal CoM KorysySitereR HNO) moor epee enie cr sca ts-nae) 1ecbosdecoctacsda:
36. Abdominis segmentum primum fascia distincte picolerate
semper ornatum, fascize margine anteriore haud aut vix
emarginato
Abdominis segmentum primum fascia sepissime fere unicolore
ornatum, fascia ipsa, quam precedentis, semper latiore, et
antice conspicue emarginata
37. Abdominis segmentum secundum nigrum, haud fasciatum
Abdominis segmentum secundum fascia colorata ornatum ......
38. Abdominis seementum primum et secundum (aut ex his unum)
fascia distincte bicolorata ornata, aut segmentum primum,
desuper aspectum, fere totum rubricolor est, margine postico
FEAR) pete coh cress sizpad sais Slee Nae See HEE Swe EL AR Ete dO Talal eae ah
Abdominis nec segmentum primum nec secundum coloribus
CUObUSmVAT eo aban, ms seeasteesacaee neeem ineeaceh apres aa tes Bee re ent
39. Anguli humerales pronoti anteriores distincte prominentes.
(Tegularum margo exterior albidus aut flavescens; tibia
posteriores ferruginez aut rufescentes, macula elongata
i lors Ehohereteh on) lone nbeemeden comaner aac nuacaantacoue ie simulator,
Anguli humerales pronoti anteriores aut fere recti aut obtusi,
had: produto. citessiige casio mere Se sanodt snide pekese tas eee Re Ean
Tegule nitidissime, perpolite ; parte interiore excepta, levis-
simee, haud) subtilissime punctate .......sscssseces+ ee. sceeeenetes
Teeule in parte exteriore subtilissime punctulate ...............
41, Clypeus Q levis, nitidus, glaber, distincte, remote punctatus ;
teeule (ex magna parte) flavescentes aut albide ; mesopleura
maculata ; tibiz intermedi et postice longe albido-maculatez,
tarsis rufescentibus.
¢ tibiis omnibus, metatarsisque intermediis et posterioribus
Supca, allie! vtec near e nae eanen cece eRe eee
Clypeus 2 conspicue minute pubescens, apice (quodam in
aspectu) elevato, grossius subdensim punctatus ; tegulze
nigricantes; mesopleura nigra; tibie posteriores albido-
flavo notate, tarsis nigrescentibus. (¢ ignotus
42. Abdominis segmentum primum, desuper visum, rubricolor,
colore hoe antice nigro-emarginato, postice anguste flayo-
marginato ; mesonotum profunde, zqualiter, sat grosse
punctatum ; clypeus (Q) antice distincte marginatus.
4.0
tricolor, sp.
43. Abdominis segmentum 2 dorsale valde fortiter a basi conico-
elevatum ; hoc, cum primo, late fasciato, fascia prima partem
segmenti (desuper aspecti) maximam occupante, antice tri-
angulariter nigro-emarginata. Macule pronotales ab angulis
posterioribus dorsalibus longe remote
Abdominis segmentum secundum haud distincte conico-
tuberculatum
44, Pronotum usque ad angulos dorsales posteriores coloratum “8
Macule pronotales a tegulis longe remoti..........................
45. Segmenta nonnulla abdominis apicalia, et dorsalia et ventralia,
nigra BE RESO ROCIO TIA HOU EE oA CE ere ARH
. Segmenta abdominalia omnia ‘colorata abies enlen Sepoe Bunnies s (aha
46. Scutellum sat convexum, quam densissime rugoso-punctatum ;
thorax post partem pronotalem niger (J); Q incognita.
zerophilus,
36
37
SEE er een Cee RE Sao a rere eee ser vulpinus Sauss.
Rife stein aide ee ate vulpinus excisus, st. n.
sp. 1.
donatus, sp. 0,
... despectus, sp. 0.
nh.
Sedo o eee ae roseotinclus, sp. 1.
44,
45
WASP-GENUS PARALASTOR. 573
Scutellum lave, parum convexum, haud densim ubique pune-
tatum, punctis interstitialibus minutis distinctis ; scutellum,
postscutellum et propodeum late aurantiaco-ornata __. placens, sp. n.
47. Clypeus antice distinctissime marginatus ; species magna et
TROVE, cesandeea es 3
48. Pronotum haud aut indistincte marginatum ad truncationem
(Picturatio abdominis segmenti 2 dorsalis pervariabilis,
nonnunquam colore rufo aut rufo-aurantiaco fasciam
apicalem tantum formante, nonnunquam totum seg-
mentum, triangulo nigro basali excepto, occupante.)
SOO PAUGADNOO Soc DUnaumoniaaooBnBenA care Eee eriurgus Sauss.
argentifrons Sm.
Abdominis seementum tertium et sequentia haud bene colorata,
marginibus apicalibus tantum obscure pallidioribus. Clypeus
3 haud marginatus, quam in precedente fortius emarginatus ;
abdominis segmentum primum multo minus fortiter trans-
versum, secundi fascia apicalis haud ant parum ampliata ;
GRINGO CTU Awe ies Veale St ah ypim ek ateN tees ite ts subobscurus, sp. n.
basim juxta inflexis
52. Thorax (mesonoto nigro aut tantum bimaculato excepto) et
Segmentum primum abdominale in aspectu dorsali ubique
DoS ORa Dp aU NGO OAS TEAR MRE me pet ce aaa MIMUS, Sp. N.
rufa aut aurantiaca, ceteris segmentis abdominalibus nigris. 53
Species aliter colorate, abdominis segmento secundo semper
CONGO CHOPIN oo cssccascoscabone: SpE OD aERtOLUAArs Dok URE OOS B ame Loe, 54.
53. Species parva, angustula, colore rubro ornata, alis hyalinis,
parte subcostali perpaullo infuscata .................. imuitator, sp. n.
Species sat robusta, aurantiaco-decorata, alis infumatis, parte
subcostali nigrescente .,
54. Species, quarum propodeum nigrum est, pronoto antice macu-
(Pronotum antice fere recte truncatum (sive levissime
concavum), angulis lateralibus distinctis, bene elevato-
marginatum ; clypeus levissime emarginatus ; abdominis
segmentum secundum densissime, grosse et conspicue
ape tsloUanal seanetet stellate a fer-velreistaepet state ee eee nautarum Sauss.
Sys)
59
RO WITe Caytitanias)) pret cks-t eres ey ye en NS eal debilis, sp. n.
' Abdominis segmentum secundum postice latissime rufo aut
aurantiaco-rufo colore fasciatum, fascia utrinque valde
ampliata, plus minus angulariter profunde (nonnunquam
profundissime) nigro-emarginata, segmentis sequentibus
coloratis
57
57.
58.
59.
61.
4 DR. R. C. L. PERKINS ON THE
Abdominis segmentum secundum, margines versus laterales,
distincte punctatum, scilicet punctis bene separatis ........ : 58
Abdominis segmenti 2 dorsalis margo posterior (desuper
inspectus) utrinque fortiter postice productus, angulis
acutis prominentibus conspicue munitus (¢) ...... eugonias, Sp. Nn.
Abdominis segmenti 2 dorsalis margo posterior utrinque dis-
tincte angulatus, angulis ipsis per paullo prominentibus (9).
microgonias, Sp. nl.
Abdominis segmentum primum basisque secundi pulchre pubes-
centia argentea utrinque ornata, propodei lateribus similiter
vestitis; fascia abdominalis segmenti secundi ad, aut paullo
ante, medium utrinque producta (q)).............-..-. argyrias, Sp. 1.
Abdomen haud pubescentia argentea ornatum; fascia ab-
dominalis segmenti 2 fere ad basim utrinqgue antice
jorovolbventay (( ocosnsanacas asd shscoqeseuasoDisaeHaraoEnen iC euclidias, sp. n.
Abdominis segmentum 2 ventrale fortissime post suleum supra
partem basalem suam abrupte elevatum, truncatione summa
antice subproducta et prominula, tuberculiformi ............... 60
Abdominis segmentum 2 ventrale post suleum transversum
distincte, sed haud fortissime, supra partem suam basalem
elevatum, truncatione summa haud subangulatim antice
PLoOductayet wUberculito nM | eee te ee ble pein ee sees eee neste 61
Segmentum abdominale tertium cum sequentibus nigrum ;
SOROS! MNES (EH) cn canconeoasdgnantoussencondadoo coos occidentalis, sp. n.
Seementum tertium et sequentia aurantiaco-ornata ; species
magnitudine mediocris (gf Q).......-.....20-e0e es .... orientalis, sp. n.
Abdominis segmentum 2 ventrale nigrum, postice aurantiaco-
marginatum, post truncationem leviter impressum (<j).
solitarius, sp. N.
Abdominis segmentum 2 ventrale colore flavo et rufo-brunneo
ornatum, nulla parte nigra, post truncationem valde late
depressum, ita ut truneatio summa incrassationem elevatam,
plerumque curvatam, prabeat (d)...............-..... hilaris, sp. n.
Thorax, pronoto excepto, niger; abdominis segmentum secun-
dum dorsale flavum, colore flayvo maculam magnam nigram
subtriangularem includente. Postscatellum tubereulatum.
maculiventris Sauss.
Thoracis pars post-pronotalis flavo-maculata ; abdominis seg-
mentum 2 nigrum, postice flavo-fasciatum, fascia nonnunquam
utrinque fortiter basim versus producta ; postscutellum
lidteperoasy he SARA A, Ot se eee el eR TT mare cia a Sth oe bicarinatus, sp. n.
. Abdomen fasciis 2 abdominalibus flavis aut sulphureis, an-
gustis, haudyuaquam dilatatis, tantum ornatum ; pronotum
antice et scutellum maculis duabus, pallide flavis, ornata.
(Amitennee U2 -articulatess))) eee eee eee eee eee optabilis, sp. n.
Picturatio cum colore valde diversa .............. Hiei aes an emis 64
, Alee atree, ubique (in quodam aspectu) violaceo-tinctee ; ab-
dominis segmentum 2 latissime fasciatum, fascia antice
angulato-emarginata. Thorax exempli unici totus niger,
postsecutello. tuberculo debili munito (Q) ........... . atripennis, sp. n.
Ale fusco-nitentes (subasneo-micantes); abdominis segmen-
tum 2 fascia apicali aurantiaca, fere simplici, ornatum ;
thorax, mesonoto excepto, ubique aurantiaco-ornatus, post-
scutello hand tuberculato (QO) 0... ee. eee teen e ees awreocinctus Guér.
. Clypeus profundissime emarginatus, emarginatione multo, quam
semicirculus, profundiore, dentem utrinque longum et ob-
WUHAN, WOMTEAAIE. ” Gorictneopnesaneen sacs Eesti AL nce ieee a, meine 5 69
Clypeus haud ita emarginatus ................. SNe cit aN ae Fe 66
Thorax (mesonoto excepto) ubique rubro-decoratus ; abdominis
segmentum primum rubrum, basali parte nigra, segmentis
ceeteris nigris........ bai decinaiea dent ee ap reat s---svene entiger, Sp, n.
Species a preecedente colore valde diverse ........................
67.
68
69.
70
Tile
72,
73.
76,
WASP-GENUS PARALASTOR.
Species aurantiaco-ornatus ; scutellum macula magna trans-
versa, vix aut haud divisa, ornatum(d) ............ infimus, Sp. Ds
Species una aurantiaco-ornata, altera colore flaviore, vix
aurantiaco, decorata; scutellum maculis duabus, late
SCOMUTTILAISL,,Onpatranany (lO) ek ae, See cee ee ora oe eee a
Fascia abdominalis prima fere simplex, lata; secunda late
emarginata, latera versus ante medium segmentum basim
versus producta ; propodeum nigrum (Q)............ sUMMUS, SP. D;
Fascia abdominalis prima antice conspicue emarginata ;
secunda fere simplex, latera versus perpaullo ampliata (Q).
mredius, sp. n.
Mesonotum solum nigrum, partibus ceeteris thoracis dorsalibus
aurantiaco - decoratis; abdcminis segmentum secundum
fascia apicali, vyix aut perpaullo latera versus dilatata,
ORTIA LUM re nC PREECE LO horn ee. ORM ORV Cn Une eustomus, Sp, n.
Thorax totus, duabus maculis pronotalibus exceptis, niger ;
abdominis Bepmontom 2 fascia latissima, antice emarginata,
utringue ante medium segmentum producta, ornatum (@).
68
pseudochromats, sp.
Clypeus profundissime emarginatus, emarginatione semicir-
culum formante, aut semicirculo profundiore ; frons capitis
OUGUITAT Lay os yn tear ee PEON gn pace ene ain ETADAS TOPS AN SEE 6) Os
Clypeus late, leviter emarginatus ; frons capitis duabus lineis
leevibus, ammetas et elevatis, conspicue munita ........ 72
Species parva, sed sat robusta ; abdominis segmentum 2 dorsale
fascia, utrinque valde anmyelhinge, ornatum ; pronotum solum
maculatum, thoracis partibus ceteris nigris (Q ),
(Clypeus colore variabilis, aut totus niger aut rufo-
AMACHIAUISS hare veeiet rycen cia aang eae
Species magnitudine mediocris; abdominis segmentum secun-
dum, cum sequentibus, nigrum ; thorax, mesonoto excepto,
TEPER OVA Mey AVIS CO) )) eaeses Se cel ARAE ona a ori ares eee Miers fallax, sp. n,
Species aurantiaco-ornata; abdominis segmentum primum
fascia lata aurantiaca, haud aut vyix emarginata, ornatum ;
scutellum, margine posteriore excepto, aurantiacum (@ ).
nN.
vulneratus Sauss,
brisbanensis, €p, ns
Species flavo-ornata; vix aurantiaca; abdominis segmentum
primum fascia flayescente, antice conspicue emarginata,
ornatum; scutellum maculis duabus, bene separatis,
Sigma human VON eur ten aay oma ee met oar Peay OB
Species maxima (14-15 mm. ad apicem seg. 2 abdominalis) ;
abdominis segmentis 2 basalibus aurantiaco-fasciatis, fasciis
haud latis, secunda simplici, hand aut vix latera versus dila-
tata, segmentis sequentibus nigris. Tibiee nigre ; posteriores
macula rufa aut flava signatee; intermedie et posteriores
(nisi attritz) pilis longis sparsim vestitee ............ princeps, sp. n,
Species aut preecedente multo minores, aut colore dissimiles ,.. 74
Segmenta 2 basalia abdominis fasciata, ceeteris nigris ; species
graciles plus minusve paralleliformes ........................--. 75
Segmenta abdominis complura aut omnia colorata; species
77
adeer act OSpane etre a centr enue ey aioe: ents eke sane ta semasle ae
Pronotum colore rufo fere ad tegulas decoratum ; clypeus 9
UO MAC Ul aye Aen eR aa erat Sher a eee eee commutatus, sp.
Pronotum aut nigrum aut maculis parvis duabus aurantiacis
ab ronSsvormatum~siclypeus O)miger ..)- 622.6 )-5s- ok. esse: 7
G abdominis segmenta 5 apicalia pilis erectis sat conspicue
vestita; tibiz utriusque sexus nigra, basi et apice nonnun-
quam pallescentibus,
(oi Ghyaswes tikyares, ©) amiensie)) 4) oaubuoccna-neceeanees suboloris, sp. n.
d abdominis segmenta apicalia pubescentia minuta depressa _
tantum vestita; tibie omnes cum tarsis rufescentes.
oloris, KP. D.
mackayensis, ep.
Nn,
n.
576 DR. R. C. L. PERKINS ON THE
le heh Claes) THENTUG) sa6.cnbocoreescocose soa sepconstss Oe ane ear a 78
© @ Clypeus totus, aut ex parte magna, niger ..... pasH a cooeads 80
78. Clypeus in parte sua media longitudinaliter impressus, utrinque
carinis subobsoletis (rotundatis nec acute eleyatis) parallelis
instructus ; antennze (ut opinor) tantum 10-articulate,
articulis 2 apicalibus minutissimis. Cellula 2 eubitalis
haud petiolata (an semper?) Segmentum abdominis ven-
traley fasab fOrbiber COMVEXUMD =o... tesa. dasisee sar = -ls odyneroides, sp. Tt.
Clypeus more solito formatus, convexus ; antennze 12-articu-
latee,fortiter clavate, articulis 4 ultimis parvis sed distinctis ;
segmentum abdominis ventrale 7 deplanatum.
(Abdominis seementum 2 dorsale fortiter elevatum pilis-
que longis erectis vestitum.) ..................64 viduus, Sp. Te
79, Abdomen fasciis duabus, hand utrinqgue ampliatis, ornatum,
prima angustissima, secunda haud lata; clypeus utrinque
longitudinaliter flavescens aut albido-flavus. Abdominis
segmentum 2 dorsale pilis erectis longioribus vestitum. {
: tuberculutus Sauss.
80. Tibis tarsique rufescentes ; cellula 2 cubitalis petiolata.
(Color variabilis ; abdominis segmentum secundum late
ferrugineo- aut aurantiaco-brunneo-fasciatum, fascia
nonnunquam basim segmenti utrinque attingente.)
carinatus Sm. (=lateritivs Sauss., var. ?}
Tibise tarsique nigrescentes, his plus minusve atro-brunneis,
articulis nonnullis testaceis ; tibize posteriores, basim versus,
in parte interiore, rufescentes. Cellula 2 cubitalis hand
petiolata (an semper ?). Segmenta abdominalia fasciis fer-
rugineis aut brunneo-aurantiacis ornata, fasciis apicalibus,
quam basales, pallidioribus.
(Nescio quomodo sint variabiles picturatio et alarum
THE UN EULOS)) Mag veeartas ae ne cee oe ava odyneripennts, sp. D.
81. Abdominis segmentum primum fascia sat lata, unicolore, rnbra,
antice angulariter nigro-emarginata, ornatum (Q)). mutabilis, sp. 1.
Abdominis segmentum primum fascia flava, albido-flava, aut :
flavo-aurantiaca anguste marginatum, aut fascia latiore,
ConSplcuel bicolonaba, Gecone tind sare: eeniey ee seeeessieeee ames 82
82. Abdominis segmentum primum fascia apicali unicolore
OLcYe(oha7h nbbns Danener ony n Oaenoe eee eb Rana aM an Gaad anoeaPincconmadee 83
Abdominis segmentum primum fascia conspicue bicolore
ornatum.
83. Mesopleura tegulas juxta seepe flavo-notata ; abdominis seg-
mentum primum transversim parum evidenter depressum :
species evidenter minor :
a. Mesopleura flavo-notata, abdominis fascia (aut prima)
pallide flavescentes aut albido-flave (Q). plebeiws, sp. n.
b. Mesopleura haud notata ; abdominis fasciz-flavee... var. a.
Mesopleura haud fiavo-notata ; abdominis segmentum primum
transversim sat conspicue depressum; species evidenter
major, fasciis abdominalibus aurantiaco-flavis, prima, quam
in specie preecedente, latiore et sat copiose punctata ( Q ).
subplebetus, sp. 1.
84. Species parva, angusta; antennsee Q fortiter ¢ fortissime
clavate, his 12-articulatis, articulis tribus apicalibus uncum
[SRRETTENA TOWNE WONT OIS, co. ocnadesaeecocsaues suuncegsoson ee debilitatus, sp. 1.
85. (Hac in sectione maris (ubi mas est cognitus) antennze tantum
10-articulate sunt, articulis duabus apicalibus minimis, intra
articulum octayum receptis.)
Abdominis segmentum primum, parte nigra excepta, unicolor,
aut flavum aut aurantiacum aut subrufum .................. We 87
Abdominis segmentum primum, parte nigra basali (si colo
niger adest) excepta, colore rufescente et flavo distincte
COLOLADUNTMI MME a emma sae mats cranes enaaue ore oases eaten : 36
86.
87.
88.
WASP-GENUS PARALASTOR. od
Clypeus $ Q lete flavus; macula capitis ocellaris Q utrinque
ad, aut fere ad antennarum insertionem utrinque producta.
nvulticolor, sp. n.
Clypeus 9 rufescens aut ferrugineus ; macula capitis ocellaris
antice emarginata, parte utrinque producta ab insertione
antennarum longe remota. G4 mihiincognitus... brwnnews Sauss.
Abdominis segmentum secundum juxta apicem transversim
impressum, ita ut incrassatio, distincte subelevata, formetur,
incrassatione postice abrupte verticali, haud obliqua, mar-
gine apicali deplanato lato. Color niger, rufo-decoratus (9 ).
anostreptus, sp. n.
Abdominis segmentum secundum juxta apicem plus minus
incrassatum, incrassatione haud antice elevata, et postice
plus minus oblique truncata; partes colorate plerumque
aurantiace aut flave aut plus minus rufo-aurantiace......... 88
Clypeus 9 aurantiaco-rufus ; oculi usque ad eclypeum colore
marginati; maculi pronotales fere ad angulos dorsales
posticos extensi; species nigra, rufo-ornata...... dyscritias, sp. 1.
Clypeus 9 niger, lateribus deflexis flavescentibus, apice rufes-
cent2; oculi haud usque ad clypeum marginati; pronotum
antice maculis triangularibus ornatum ; species nigra, flavo-
OIA IEDT As vanontas abies darter OBOSe OHO Mees cando aud saben wanthochromus, sp. 1.
Divisio secunda.
Thorax et abdomen bicolorata, nigra et rufa sive ferruginea ... 14
Corpus nonnulla in parte semper flavo aut albido aut auran-
tiaco-flavo colore ornatum, spe tricoloratum ..................
Teeule (parte interiori thoracem juxta excepta) ex magna
parte polite, impunctate aut fere impunctate, sculptura
nulla aut vix videnda, puncta rare hic illic disposita non-
TU TENA TELE TILE Supra ener eee GEV ee erst steal stettele ae res 3
Tegulze plerumque subtilissime sed distinctissime in parte
exteriori punctate, nonnunquam sat copiose et grosse
12)
punctate, aut puncta grossa et minuta intermixta ferentes... 8
Abdominis segmentum primum supra rubens, margine postico
LEWTO 2 [OOO IAM TUK 55 coc sey coasoucdvoomece picteti Sauss.
Abdominis segmentum primum aliter coloratum; propodeo
DY Coloration setters as t-ceeet ater ts ace pemeiermcte ne seme ere tea A
Abdominis segmentum secundum postice fascia flavo-aurantiaca
lata ornatum, partem, majorem quam quartam, segmenti
occupante, ad basim utrinque conspicue aurantiaco-
maculatum.
(Clypeus truncatus aut parum emarginatus.). constrictius, sp. n.
Abdominis segmentum secundum nonnunguam haud fasciatum,
aut fascia pallide flava, aut obscuricolore ornatum ; si aur-
antiaco-fasciatum, fascia minus lata est, haud quartam
segmenti partem occupans.
Color flavus superficiem totam posteriorem segmenti primi
ocecupans, colore nigro, si adest, superficiem declivem solam
OC CUP ATC Mees ie -1- Say ao ETT. SRE PRE rs eee 6
Abdominis segmentum primum fascia apicali tantum ornatum. 7
Tegule Q haud flavo-maculatx, feminz et maris punctis non-
nullis grossioribus et profundioribus signatee ... mesochlorus, sp. n.
Tegule femine flavo-maculate, feminze et maris punctis minus
grossis et profundis signatze ......... mesochlorws mesochloroides, st. N.
Abdominis segmentum 2 ventrale medium haud impressum ;
Q clypei apex late leviterque emarginatus ; mesonotum
duabus lineis elongatis subparallelis ornatum; 3 incognitus.
(Abdominis color atro-brunneus, segmentis fasciis albido-
LIENS OMAN AMISY)) Danacomad adckad .geboctap ae qonnoAtn sods . darwinianus, sp. D.
578 DR. R. C. L. PERKINS ON THE
Abdominis segmentum 2 ventrale post sulcum transversum
conspicue impressum, ita ut truneatio summa rotundatim
elevatur (¢).
(a) Thorax nigrum, flavo-notatum ............... comptus, sp. 1.
(b) Pronotum rufum, flayo-notatum, mesonoto medio rufo.
comptus, var. rwbescens, 1.
8. Abdomen brunneum, rufo-brunneum aut atro-brunneum, et
colore albido aut flavo-albido variegatum ......................-. uy
Abdomen nigrum, colore sulphureo, flavo aut aurantiaco
VLELE Deu bl pe yop eney yon. Ae ATU. coc s a SbUdide wistote ells asian oie 10
9. Tegulz subtilissime ubique punctate ; puncta nonnulla majora
sed levissime impressa adsunt; abdominis segmentum 2 dor-
sale maculis basalibus carens. aut his minimis...... alecandriz, sp. n.
Tegule subtilissime punctate et etiam punctis grossioribus,
et, quam in precedente, majis conspicuis, signatze ; abdominis
segmentum 2 dorsale maculis duabus, conspicuis, basalibus
OUTA DDI erat ety ahah oee cs ac nvacelaeee nas ace arenicola, sp. n.
10. Clypeus truncatus aut vix emarginatus ; ¢ tegule fere ubique
grosse et conspicue punctate. 9 imcognita ...... simillimus, sp. Tl.
Clypeus distincte, sed leviter, emarginatus; tegule minu-
tissime copiose punctate, punctis grossioribus inter punetula
Minuta NOMDUNGUAM IMNGETSPETSIS ............... eee cece eee ee ee inl
11. Abdominis seementum 2 basalinm fasciz evidenter bicolores,
antice aut aurantiace aut rufescentes, ilies flavescentes
EEN ] OSVE(CONMNIES Sam anpoaroaes soo scobceasanenondsee ; 12
Abdominis seementum secundum plerumque nigrum, aut postice
tantum piceo-marginatum, aut fasciam incompletam habens ;
primum in aspectu dorsali totum (aut fere totum) flavum, aut
yoMMOHE CET, IEHOUSL loRKeoMKO MARTIN oO ok apd ghennonnenn cre ron ook 13
‘12. Femora posteriora ubique aut ex magna parte fulvescentia ;
Pee facie sua et forma cum finitimis congruens:
a. Stigma lucide flavescens ...... vce Synchromus, sp. 1.
b. Stigma obscure flavescens aut aoe venice tte RS
Femora posteriora nigricantia, apicibus plus minus rufescenti-
bus; species elongata, angustissima, abdominis segmento
secundo plus, quam solito, elongato et angusto.
(Stigma atro-brunneum aut nigrescens.) ........._ leptias, sp. n.
13. Pronotum ¢ totum flavum aut subaurantiacum ; alarum stigma
atrum.
(Abdominis segmentum primum supra aurantiaco-flaypm.
secundum fascia imperfecta apicali, duabusque maculis
parvis subbasalibus, vix conspicuis, ornatum; anten-
narum flagellum nigrum, apice extremo rufescente.
Picturatio segmenti 2 abdominis an constans ?) ¢.
tgnotus, Sp. NT.
Pronotum g et Q haud totum flavum, nonnunquam antice
flavo-notatum ; alarum stigma lucide flayum.
(Abdominis segmentum primum supra flavum, secundum
nonnunquam totum nigrum aut margine apicali piceo
aut rufescente ; pronotum colore variabilis, supra rufum
aut nigrum, aut rufo-marginatum, nonnunquam antice
ornatum maculis duabus distinctis flavis.) ... icamotdes, sp. D.
14, Tegule ubique copiose, subtilissime punctate ; species major.
mimus.
Tegulz conspicue grosse punctate, inter puncta nitidee, pune-
tulis minutis, vix aut haud diseernendis ..........., imitatoy.
WASP-GENUS PARALASTOR. 579
1. Paratasror (Paralastoroides) cLorHo Lep.
Distinct from all other species by the suture of the first
abdominal segment and by the coloration. I have not seen this
species, and J do not not know whether Saussure saw more than
a single example. He says that the suture is ‘ more or less
distinct.”
Hab. Australia.
2. PARALASTOR SAUSSUREI, sp. 0.
2. Nigra, fronte interantennali rufo-punctata, capitis vertice
utringque minute flavo-notato; abdominis segmento secundo,
triangulo elongato nigro mediano excepto, cum sequentibus, ferru-
gineo, aut plus minus aurantiaco. Clypeus nitidus, haud densim,
sed plus minus strigose, punctatus, apice distincte, levius
emarginato. Frons capitis grosse et distincte punctata, linea
leevi, semicirculari, ocellum anteriorem includente, instructa.
Capitis vertex et thorax densissime rugoso-punctati, postscutello
medio elevatulo, sive tuberculato, et, cum propodeo, opaco et
multo, quam scutellum, subtilius sculpturato. Tegule fortiter
conspicue punctate. Abdominis segmentum primum, pars
secundi dimidia basalis, partes frontis capitis orbitales, et basis
et latera deflexa clypei, albido-tomentosa, partibus abdominis
ceteris aureo-tomentosis. Al superiores fortiter infuscate, et
cexruleo-nitentes. Segmentum abdominis 2 ventrale post suleum
valde fortiter elevatum, elevatione summa prominente. Long.
a fronte usque ad apicem segmenti 2 abdominis 15 mm.
Hab. North Queensland (Dodd).
3. PARALASTOR INFERNALIS Sauss.
The band of the second abdominal segment varies in colour
from pale orange-yellow to deeper orange, and is sometimes
angulately sometimes roundly emarginate. The facial markings
of the male vary, the clypeus sometimes being wholly yellow,
sometimes only yellow basally, with intermediate conditions.
The sinus of the eyes is sometimes yellow, the scape in front and
a medio-frontal spot probably always so. The pronotum is
usually spotted with yellow, more rarely black. Three apical
joints of the antenne small, but distinct. Tegule with
conspicuous, deep, coarse punctures.
Hab. Cairns (3 (N. Queensland) and Port Darwin. Twenty-four
exainples examined.
4, PARALASTOR RUFIPES, Sp. Nn.
Agrees generally in form and in sculpture with P. infernalis,
but is at once distinguished by the red legs and the narrower
apical band of the second abdominal segment, which is only a
little dilated at the extreme sides. The postscutellum is without
the distinct tubercle of infernalis 9, in which species it may be
580. DR. R. C. L. PERKINS ON THE
obscure in the male, but is well developed in the other sex. Size
of infernalis.
Hab. North Queensland (Dodd), 1 2.
5. PARALASTOR FRATERNUS Sauss.
The male has the clypeus (at least for the most part), the frontal
spot, and the front of the scape of the antenne orange or
yellowish orange. The tegule are for the most part very minutely
punctured and clothed with very short hairs or tomentum.
Hab. The single male in the Oxford Museum is from New
South Wales, the female type in the British Museum has no
special locality assigned to it.
6. PARALASTOR CONSPICUUS, Sp. N.
3. Niger; clypeus, macula elongata frontalis huie adjuncta,
articulus antennarum primus antice, maculeque 2 pronotales
pallide flava, aut albido-flavescentia. Abdominis segmentum
secundum fascia lata pallida basali, postice profunde triangulariter
emarginata, et antice plus minus nigro-emarginata, ornatum,
Clypeus profundissime emarginatus. Frons capitis ante ocellum
anteriorem grosse punctata, punctis inter se distinctis. Anguli
pronotales prominuli. Mesonotum sat grosse et distincte punc-
tatum, exemplorum recentium dense nigro-tomentosum, pilisque
erectis breviorilbus subpallidis vestitum. Scutellum tuberculo
prominente munitum, propodeo antice grosse punctato. Tegule
dense et minutissime ex parte majore punctate, punctis magnis
nonnullis presentibus. Abdominis segmentum primum fortiter
crebre punctatum, punctis sepe sub tomento denso abditis ;
segmentum 2 ventrale post suleum valde fortiter elevatum,
elevatione summa prominente; segmentum 7 ventrale_pilis
erectis minus brevibus vestitum. Antennarum articuli 3 ultimi
minimi., Al, costali parte excepta, hyaline. Long. 11-12 mm.
Hab. North Queensland, Cairns district (Dodd). Vive
examples.
The sculpture of the tegule, as in fraternus, is quite different
from that of P. imfernalis and its allies, the coarse punctures
being altogether confined to their inner half (or almost so), the
outer part bearing the very fine puncturation.
7. PARALASTOR DUBIOSUS, sp. n.
3. Hardly differs from P. conspicua in structure, and is pro-
bably only a race of that species. The second abdominal segment
is entirely pale, or has a more or less extensive median longitudinal
dark marking, The silvery tomentum, that fills the ocular sinus
and extends down over the deflexed sides of the clypeus, is more
dense and conspicuous and the sculpture of the latter beneath it is
somewhat changed in accordance.
This form varies much in detail of colouring, as shown partly
in the table of species above. In one example the pronotal spots
WASP-GENUS PARALASTOR. 581
are extremely small, while in another they form a broad band,
slightly interrupted in the middle.
Hab. Queensland, Mackay (Zurner), 5 3; Queensland, from
E. Saunders’s collection, 2 3.
8. PARALASTOR COGNATUS Sm.
This species is entirely distinct from and not closely allied to
P. fraternus, which it resembles in colour, and of which Smith
thought it might be the male. Its comparatively slightly
emarginate, bicarinate clypeus distinguishes it at once.
Hab. Doro (Wallace); type in the Oxford Museum.
9. PARALASTOR UNIFASCIATUS Sm.
Clypeus formed as in P. cognatus; the tegule shining, with
the minute puncturation not, or barely at all, discernible, whereas
in cognatus their surface is nearly dull, and there is a copious
though excessively fine puncturation,
Hab. Aru (Wallace) ; type in the Oxford Museum.
10. PARALASTOR CONSPICIENDUS, Sp. n.
@. Nigra; clypei pars basalis et macula interantennalis,
nonnunguam sinus ocularis, macula magna utrinque postorbitalis,
pronoti margo anterior late, fasciaque lata basalis segmenti
secundi abdominalis, postice emarginata, flava aut albido-flava.
Clypeus late lunulato-emarginatus, utrinque carina, bene elevata,
acute munitus. Frons capitis. dense aureo-tomentosa, distincte
fortiter punctatus, tomento remoto. Anguli pronotales distincti,
haud prominuli, Mesonotum tomentosum, et pilis erectis sat
evebre vestitum, postscutello haud evidenter tuberculato. Tegule,
marginem exteriorem versus, minutissime punctulate, ibique
haud aliter punctate. Ale, costali parte atro-fusca excepta, fere
hyaline. Abdominis segmentum 2 ventrale fortissime abrupte
supra suleum elevatum, elevatione summa antice prominula.
Long. 12-13 mm.
Hab. Queensland, Inkerman, near Townsville (Stalker), 1 9 ;
Townsville, 1 2 (Dodd); Queensland, 1 92, from EH. Saunders.
All in the British Museum Collection.
11. PARALASTOR ELEGANS, Sp. n.
2. Colore et picturatione P. maculiventris Sauss. (Hit. Masar.
et Suppl. pl. xvi. f. 3).
This species differs from Saussure’s description of P. maculi-
ventris only as follows :—The clypeal emargination between
the teeth is not straight, but slightly rounded; there is only a
yellow spot on the inner orbits, the head is very densely clothed
with golden tomentum, not blackish. The fascia of the first
abdominal segment is incomplete, failing before the lateral
angles.
582 DR. R. C. L. PERKINS ON THE
The clypeus has two very strongly raised longitudinal carine,
not mentioned by Saussure, but which, as Mr. Meade-W aldo has
kindly informed me, are present in the type, the face of which is
asymmetrical or somewhat deformed. In most respects P. elegans
resembles P. conspiciendus, apart from the quite different colour-
pattern, but the clypeus is much shorter and the propodeum
shorter and more rounded, less quadrate.
Hab. N. Queensland (#7. P. Dodd), 2 2.
12. PARALASTOR TRICARINULATUS, Sp. 0.
3. Niger, clypeo, maculaque elongata interantennali, huie
adjuncta, flavis, macula parva verticis postoculari utrinque
aurantiaca. Abdominis segmentum primum spatio lunulato
apicali rufo-decoratum, secundum ferrugineo-rufum, macula
basali, haud ad medium segmentum extensa, nigra, segmentis
sequentibus cum tibiis tarsisque aurantiacis aut ferrugineis.
Clypeus leviter emarginatus, apice lato, anguste _nigro-
marginato, evidenter, sed parum _ acute, longitudinaliter
3-carinatus. Frons capitis rugoso-punctata. Pronoti truncatio
vix marginata. Mesonotum grosse et rugose punctatum.
Scutellum postice conspicue foveatum. Postscutellum medium.
subproductum sive subtuberculatum. Abdominis segmentum
primum transversim subconspicue impressum, puncturatione sua,
quam mesonotali, minus grossa; segmentum 2 ventrale post
sulcam abrupte fortiter elevatum, elevatione summa _paullo
rotundatim prominula, grossim et copiose punctatum ; segmentim
7 ventrale pilis erectis sat longis conspicue vestitum, medium
longitudinaliter sulcatum. Ale, costali parte excepta, subhyaline.
Long. 13-14 mm.
Hab. Victoria (C. French), 1 3.
13. PARALASTOR TASMANIENSIS Sauss.
This species was described by Saussure from a headless male,
and is said by him to inhabit Tasmania, but I have only seen
Queensland examples.
In the male the clypeus, medio-frontal spot, and front of the
scape of the antenne and the postocular spots of the vertex are
yellow, the latter sometimes redder or orange. In the female
the apical portion of the clypeus is black, the rest orange or fer-
ruginous brown. The scape of the antenne black or largely
ferruginous or reddish brown in front. The tegule are very
coarsely punctured, the punctures extending to (or almost to)
the outer margin. The apical ventral segment of the male is
wide and widely impressed, and bears only very short hairs, as
do the preceding segments.
Hab. Queensland, Brisbane; in Oct., Jan., Dec., and March
(Hacker). I suspect that the locality (Tasmania) given by Saus-
sure 1s erroneous.
WASP-GENUS PARALASTOR. 583
14, PARALASTOR HABILIS, sp. n.
Abdomen nigrum, segmento secundo pallide flavo, macula,
magna subtriangulari nigra, ad aut post medium segmentum
extensa, signato. o clypeus, macula interantennalis, articulus
antennarum primus antice, macule parve verticis postoculares,
macule pronotales (nonnunquam fasciam fere integram
formantes), pallide flava; @ clypeus antice niger, postice
aurantiacus ; antennarum articulus primus (basi extrema pallida
excepta) niger. Ale hyaline, costali parte infuscata, plus minus
flavescente. _Tarsi posteriores, articulo ultimo excepto, nigri
aut atro-fusci ; tibie aut nigricantes aut rufescentes.
Clypeus distincte, nee profunde emarginatus; frons grosse
punctata. Mesonotum, quam densissime, rugoso-punctatum,
parum dense pilosum; tegulis nigris, grosse usque ad marginem
exteriorem punctatis ; posteutellum haud aut vix tuberculatum.
Abdominis segmentum primum pernigrum, dense punctatum,
inter puncta quasi subtilissime granulatum, secundi pars nigra
similiter sculpturata, sed remote punctata ; segmentum 2 ventrale
post suleum fortissime abrupte elevatum, elevatione summa
prominula; septimum segmentum ¢ ventrale latum, lateque
impressum, parum conspicue pilosum, pubescentia brevi vestitum.
Long. 11-13 mm.
Hab. North Queensland (Dodd); 2 5,1 @.
15. PARALASTOR SUBHABILIS, Sp. 0.
Precedenti simillimus, ut apparet, paullo gracilior, alarum parte
costali magis flavescente, et ibidem venis pallidioribus. Tarsi
posteriores ubique, aut ex majore parte, pallidi. Tegule,
marginem exteriorem versus, pallide. Segmentum tertium
abdominale plerumque flavo-marginatum, aut plus minusve
pallidum, rare nigricans, sequentibus segmentis etiam spe
pallidis. .
Possibly only a race of the preceding, but superficially quite
distinct. The pattern of the second abdominal segment is different
from that of Aabilis, the black median marking narrower and not
of the same subtriangular form. Unless it has become changed
post mortem, the ground-colour is also different, the yellow more
ochreous, or in some examples approaching orange, not pale clear
yellow or whitish yellow. The colour of the clypeus in the female
is not constant ; sometimes it is nearly entirely yellow, sometimes
black on the apical part.
Hab. Queensland, Mackay, 6 examples (Zurner) ; Queensland,
1 S$ from EK. Saunders’s collection.
16. PARALASTOR FLAVICEPS Sauss.
I have not examined the type of this species, but should the
tegule have a very coarse puncturation extending to the outer
margin, then it would be placed in my table next to habilis and
584 DR. R. CG. L. PERKINS ON THE
subhabilis, being easily distinguised by its “ téte jaune de soufre ;
sur le vertex un ovale noir, qui enveloppe les ocelles” (Saussure).
In any ease this character of coloration will distinguish it from
any other of the species that have the first abdominal segment
wholly black.
Hab. Australia.
17. PARALASTOR ALBIFRONS Fabr.
I have not examined the type of this species, but Mr. Meade-
Waldo has very kindly examined it for me, as to certain
structural points, from which I infer it should be placed next to
P. habilis. In fact, I do not feel sure that the latter may not be
a variety or race of the Fabrician species.
Hab. Australia (Banks collection).
18. PARALASTOR APICATUS Sin.
Tbe type specimen is from Aru, Examples from Papua
(Odynerus lorentzi Cam.) are at the most slight colour-varieties,
quite unworthy of a special name. The tegule are punctured on
their inner portion, smooth and polished outwardly ; the second
abdominal segment seen in profile rises up strongly above the
level of the first; ventrally, it rises up strongly, but rather
obliquely, from the transverse suleus, and the truncation is not at
all prominent or produced forwards at its highest point.
Hab. Aru (Wallace), type in the British Museum and 1 ¢ in
the Oxford Museum. I have seen several examples from New
Guinea.
19. PARALASTOR PALLIDUS, Sp. n.
2. Nigra, clypeo flavo-bimaculato, aut macula curvata basali
ornato. Frons interantennalis flavo-maculata ; macule pronotales
flavescentes. Tegule externe flavescentes aut rufo-flave.
Abdominis segmentum secundum ochraceo-flavum, macula
triangulari basali signatum, sequentibus similiter pallide coloratis,
margine etiam primi postico pallide fasciato, fascia hac media
emarginata. Femorum apices, tibie, tarsique rufescentia. Ale
subhyaline, parte costali infuscata. Clypeus nitidus, iregulariter
punctatus, apice distincte, nec profunde, emarginato, margine
elevato. Mesonotum densissime punctatum, postscutello inermi.
Abdominis segmentum primum obscurius punctatum, secundum
ventrale fortissime post sulcum abrupte elevatum, elevatione
summa prominente. Tegule ex magna parte dense distinctissime
et subtilissime punctate.
Unlike any other species superficially. The second dorsal
segment is only a little more than ordinarily convex on its basal
portion.
Hab. Queensland (E. Saunders’s collection), 1 9 ; Mackay
(Turner), 1 &.
>
WASP-GENUS PARALASTOR. 58
>) |
20. PARALASTOR INSULARIS Sauss.
Unlike any other species superficially, the second abdominal
segment black, with the extreme apex only indistinctly pale.
Hab. Australia, Swan River; 1 ¢.
21. PARALASTOR TUBERCULATUS Sauss. (PI. I. fig. 3.)
A very distinct species, of which I have seen six females, four in
the British Museum Collection and two in the Oxford Museum.
This species should be removed from this position and placed
next to P. odyneroides (no. 67).
Hab. Tasmania; Adelaide: Victoria (Yrench). In the latter
locality the abdominal bands are deeper yellow, in one example
more orange-colourcd.
22. PARALASTOR EMARGINATUS Sauss.
Hab. Tasmania, 1 @ in the Oxford Museum; Eaglebawk
Neck, 5.H. Tasmania, Feb. 12th—March 3rd, 1913,1 9 (Zurner).
23. PARALASTOR PARCA Sauss. (PI. I. fig. 1.)
In one or two females the clypeus is quite truncate at the apex,
and others are intermediate between these and those in which
it is quite distinctly, though always shallowly, emarginate.
Hab. Tasmania, Franklin; Mt. Wellington and Eaglehawk
Neck (Turner); Victoria (French).
24, PARALASTOR LETUS, sp. n.
3. Niger, clypeo, antennarum articulo primo antice, macula
interantennali, duabusque postocularibus, tlavis. Pronotum,
scutellum et nonnunquam postscutellum, flavo-bimaculata.
Mesopleura sub tegulis flavo-notata, his ex magna parte rufes-
centibus, et nonnunquam flavo-notatis, Abdominis segmenta
2 basalia fascia flava apicali ornata, fascia prima media levius
emarginata, secunda simplici, haud lata. ‘Tibie tarsique
rufescentes. Ale distincte infuscate. Clypeus distincte, nec
profunde, emarginatus, pilis pallidis ubique vestitus. Frons
capitis dense rugoso-punctata, capite pilis longis vestito.
Mesonotum dense punctatum, postscutello mermi; propodei
lateribus rotundatis. Abdominis segmentum primum fortiter
transversum, parte apicali excepta rugosissime puncturatum ;
segmentum secundum dorsale basim versus mediocriter convexum,
pilis erectis longioribus parce vestitum; secundum ventrale
fortissime et abrupte post suleum elevatum, elevatione summa
prominente ; segmentum apicale ventrale pilis erectis sat longis
vestitum, haud distincte depressum. Long. 9-10 mm.
Hab. Fremantle, Australia (British Museum), 2 ¢.
25. PARALASTOR FRATER, Sp. n.
3. Niger, capite thoraceque ut in P. lato, maculatis.
Abdominis fascia prima et secunda fere wqualiter lati, simplices ;
586 DR. R. C. L. PERKINS ON THE
postscutellum nigrum. A P.leto segmento tertio dorsali cum
sequentibus pilis nullis erectis vestito, segmento 7 ventrali, cum
preecedentibus, pilis erectis tantum brevissimis vestito, facile
distinguendus. Tegule minutissime punctate. Abdominis
segmentum primum latissimum et cum secundo nigerrimum.
Certainly distinct from the preceding by the differently clothed
apical male ventral segment—a very important character. In
both the tegule are for the most part feebly and very minutely
punctate, the punctures feebly impressed.
Hab. Albany (Brewer); 1 3 in the Oxford Museum.
26. PARALASTOR ORDINARIUS, Sp. n.
9. Nigra, clypeo albido-flavo, macula diseali nigra; macula
interantennalis, una utrinque postocularis, et due pronotales,
pallide flave. Tegule nigricantes, in parte exteriore testacesx,
postice flavo-notatee. Abdominis segmentum primum postice
albido-flavo-fasciatum, fascia utrinque paullo dilatata ; segmenta
sequentia simpliciter (nec late) fasciata. Tibi tarsique
rufescentes. Clypeus latus, brevior, apice late et levissime
emarginatus. Frons capitis crebre punctata. Mesonotum dense,
minus grosse punctatum, postscutello inermi. Propodei latera
fere equaliter rotundata. Tegule ex parte magna lzevissime,
polite, impunctate, Abdominis segmentum primum subcrebre
punctatum, fascia pallida apicali puncta multa ferente. Ab-
dominis segmentum secundum dorsale basim versus subfortiter
convexum, ventrale, post suleum, fortiter elevatum, elevatione
summa haud prominente. Ale subhyaline, parte costali magis
infuscata, stigmate medio pallescente. Long. 85mm.
A rather commonplace species, chiefly noticeable as having
several pale abdominal bands.
Hab. Victoria, Wimmera (coll. Froggatt), 1 2.
27. PARALASTOR PUNCTULATUS Sauss. (PI. I. fig. 17.)
(3 =P. albocinctus Sm.% 2, var. =similis Sauss. 2)
In this species the apex of the clypeus is not always quite
truncate, but is sometimes very faintly concave or has the margin
a little sinuate. The basal abdominal segment is less strongly
transverse than in many of the allied species. It is certainly
variable both in small details of colour and structure, and I
believe that P. albocinctus Sm. is only a male of this species, and
similis Sauss. a slight variety. Saussure, in his first deseription
of the latter, says that the second ventral segment of the abdomen
is “sans tubercule,” and in his supplement he gives the presence
or absence of a tubercle as the only distinctive character between
the two forms. As in other species I have found that the trun-
cation of the segment is sometimes more evidently raised or
produced in some examples than in others, this minute distinction
seems hardly sufficient.
The male of punctulatus is abundantly distinct from any of
WASP-GENUS PARALASTOR. 587
the similar forms described by Saussure by its 12-jointed (not
11-jointed) antenne.
Hab. Tasmania, Mt. Wellington and Eaglehawk Neck, Jan.,
Feb., and March, 1913 (Turner); Hobart; Mt. Kosciusko,
6000 ft., N.S.W. (Waterhouse).
28. PARALASTOR OPTABILIS*, sp.n. (PI. I. fig. 12.)
Picturatio capitis, thoracis et abdominis, cum P. fratris et
P. leti picturatione congruens. Species robusta, alis infuseatis.
Clypeus 9 medius niger. Clypeus distincte dentato-emarginatus,
margine apical inter dentes laterales medio leviter rotundato, sive
paullo producto, quasi tridentato. Caput cum thorace sat dense
pilis longis vestitum. Anguli pronotales parum distincti. Meso-
notum tomentosum, punctis sepe plus minus tomento obscuratis,
postice haud dense (sc. irregulariter) punctatum, postscutello
rotundato, imermi. Abdominis segmenta 2 basalia tomento
nigerrima, puncturatione spe plus minus abdita, sive obscurata ;
primum fortissime transversum, secundum pilis erectis crebre
ubique vestitum; fascize abdominales minus late, fere recte.
Abdominis segmentum 2 ventrale fortiter abrupte post suleum
elevatum, elevatione summa prominente; segmentum ventrale
3 apicale, eque ac precedentia, perconspicue et dense pilis
erectis vestitum. Antenne g 12-articulate. Tibi tarsique
rufescentes aut testacei, plus minusve (presertim in ¢ ) flavo-
notati. Femora anteriora et imtermedia ¢ flavo-notata, his
in © etiam flavo-notatis. Long. 12(¢ )-14(2) mm.
Hab. North Queensland (Dodd).
29. PARALASTOR VULPINUS Sauss.
In its typical form this species is distinct to the naked eye by
its narrow first abdominal fascia, which, like the second. is pale
yellow posteriorly and orange- _brown or brown in front, and its
robust form—the latter ehenacien distinguishing it from other
species which have similarly bicolorous fasciz. The first abdominal
band becomes narrower towards the sides.
Other specimens often show little or no trace of the bicoloration
of the fascie, which are often orange; the first is then excised
or emarginate in the middle, so that it is notably widened on
either side. This form may, with larger and better material,
prove to be distinct; but I can see no satisfactory structural
difference. It may be known as P. vulpinus st. excisus, n.
The well-clothed second dorsal segment of the abdomen, in well-
preserved examples, is characteristic of both forms. The apical
ventral segment of the male, if fully exserted, bears at its base a
remarkable tuft of dense erect hairs, but these may be withdiawn
beneath the sixth segment. Similar tufts oceur in P. simulator,
which is evidently allied to vulpinus. The antenne of the male
are 11]-jointed.
* This species should be placed next to P. awreocinctus and atripennis, as in the
table.
Proc. Zoou. Soc.—1914, No. XLI. A]
588 DR. R. C. L. PERKINS ON THE
I have seen seven examples of P. vulpinus and ten of the race
excisus ; most of them are in bad condition.
Hab. Typical form: Adelaide, 1 2, Oxford Museum, and
1 2, British Museum; Victoria (French), 1 9 ; Croydon, 1 d
(Froggatt); the rest without special locality. Race excisus:
Pt. Stephen, 1 small 2 ; Melbourne (French), 1 2 ; Mittagong,
N.S.W., 1 2 ; Cumberland, N.S.W. (Zurner), 1 2 ; Woodford,
GOK
39, PARALASTOR SIMULATOR, Sp. n.
Picturatio thoracis et abdominis picturationi P. vulpini fere
similis; mesopleura seepissime nigra, innotata; scutellum non-
nunquam bimaculatum; clypeus 9 macula magna curvata
basali ornatus, aut hac bipartita. Tibiz tarsique ferruginel ;
tibie posteriores linea albida ornate: ale hyaline costali parte
sola distincte infuscata. Tegule nitide aut subnitide, flavo-
marginate, aut albido-marginate, plus minusve_ subtilissime
punctate. Clypeus leviter aut levissime emarginatus, 9 nitidus,
parcissime distincte punctatus, inter puncta hee majora sub-
tilissime distincte punctulatus. Frons capitis dense punctata.
Angul pronotales distincte prominentes. Caput cum thorace
puis crebre vestitum. Mesonotum et scutellum crebre, plus
minus grosse, punctata, scutello inermi. Abdominis segmentum
primum fortissime transversum; secundum dorsale (precipue
maris) fortiter supra primum convexim elevatum, ventrale
fortiter post suleum elevatum, elevatione obtusa, parum pro-
minente; segmenta sequentia ventralia ¢ brevissime pubescentia.
Antenne ¢ 11]-articulate, articulis 3 ultimis minutissimis. Long.
6°5-9'5 mm.
Hab. Adelaide, 2 ¢ ; Victoria (french), 1 9; 2 g andl 9
without special locality.
31. PARALASTOR PUSILLUS Sauss.
This species is of the same general appearance and structure as
P. simulator, but should easily be distinguished by the prothoracic
angles being indistinct or not at all produced and the second
ventral seginent having the top of the truncation in the middle
produced into a spiniform tubercle. No mention is made of a
white or yellow margin to the tegule, nor of the conspicuous
white or yellow line on the hind tibiz.
The examples that I have. before me from the British Museum
and Oxford, named pusillus, are partly P. simulator and partly
other species.
Hab. “ Lia Nouvelle-Galles du Sud ” (coll. F. Smith).
32. PARALASTOR DONATUS, sp. n.
Colore P. simulatori fere assimilis. Abdomen fasciis duabus
bicoloratis ornatum, secunda latera versus ampliata. Clypeus
WASP-GENUS PARALASTOR. 589
$ albido-flavus, 2 niger, macula permagna, curvata, basali,
aurantiaco-tincta, ornatus. 3 antennarum articuli 2 primi
antice, maculaque interantennalis albido-flavescentes; 2 arti-
culus primus antennalis antice aurantiacus, macula inter-
antennali bicolore. Macule postoculares verticis minute,
flavescentes. Macule pronotales sat magne, aut bicolores, aut
rufescentes. Scutellum bimaculatum; mesopleura aut flavo-
aut aurantiaco-maculata. Propodeum utrinque juxta basim
suam notatum. ‘Tibiz omnes ¢ cum metatarsis intermediis
et posterioribus fere tote albide; 2 tibize intermedia et
posteriores supra longe albescentibus ; femora posteriora utrius-
que sexus ferruginea; anteriora et intermedia plus minusve
albido-variegata. Tegule albido-cincte. Clypeus levissime
emarginatus, ¢ opacus, convexus, dense argenteo-pubescens,
olsoletim punctatus, inter hec puncta majora densissime et
subtilissime punctulatus ; 2 pernitidus, distincte sparsim punc-
tatus, inter hee puncta parcissime, vix evidenter, minutissime
punctulatus. Frons capitis parum profunde punctata. Anguli
pronotales fere recti, haud prominentes. Scutellum distincte,
nec dense, punctatum, postscutello inermi; propodei lateribus
rotundatis. Abdominis segmentum secundum dorsale haud
fortiter convexim elevatum, tomentosus, pilis erectis carens;
ventrale, post sulcum transversum, sat fortiter, suboblique supra
partem basalem suam elevatum, elevatione summa haud pro-
minente. Segmentum < ventrale 7 pilis brevissimis erectis
vestitum. Ale hyaline, costali parte sola infuscata; .tegulis
glaberrimis, nitidis, ex majore parte impunctatis. Long. 8—
9 mm.
Hab. Bacchus (or Bocchus?) Marsh; given to me by my late
friend, G. W. Kirkaldy.
33. PARALASTOR DESPECTUS, Sp. n.
2. Nigra, parte basali clypei macula, antice emarginata,
aurantiaca ornata. Frons interantennalis aurantiaco-notata.
Antennarum articulus primus antice flavo-lineatus. Macule
postorbitales verticis minime, flave. Pronotum rufescens,
antice utrinque plus minusve flavescens, nigro-marginatum.
Postscutellum utrinque macula parva aurantiaca aut rufescente
ornatum. Abdominis segmentum primum, parte nigra declivi
excepta, fere totum rufescens et postice albido-flavo-marginatum ;
secundum fascia simili decoratum, antice emarginata, latera
versus fere ad medium segmentum extensa. Clypeus levissime
emarginatus, margine apicali elevato, sat dense, grossius punc-
tatum, et cum oculorum sinu conspicue albo-pubescens. Anguli
pronotales haud prominentes. Mesonotum cum scutello nitidum,
illo Gense et grosse punctato, postscutello inermi. Tegule ex
majore parte glabree, polite, nigra, margine ipso testaceo. Ale
fere hyalinz, costa magis infuscata. Abdominis segmentum 2
dorsale ad basim fortiter convexum ; ventrale post sulenm fortiter
4\*
590 DR. R. C. L. PERKINS ON THE
supra partem suam basalem elevatum, elevatione summa hand
prominente sive tuberculiformi. Tibie ferruginee, intermedize
et posteriores intra infuscate sive nigricantes, his supra flavo-
lineatis, tarsis nigricantibus, articulo apicali pallescente. Antenne
breves et crasse. Long. 8 mm.
Hab. W. Australia; 1 2 in the British Museum.
34. PARALASTOR TRICOLOR, sp.n. (PI. I. fig. 5.)
2 colore P. despecto simillima, sed postscutello haud binotato,
scutello bimaculato, mesopleuris notatis, abdominis segmento
secundo minus late fasciate, fascia fere recta.
3 clypeus flavus aut flavus et aurantiacus, medius fusco- aut
nigro-maculatus, Antennarum articulus primus rufescens,
antice flavescens, secundus nonununguam rufescens. Macula
interantennalis, macule pronotales, scutellares et mesopleurales
aurantiace, aut plus minus flavescentes aut rufescentes. Tegule
rufescentes aut flavo-variegate. Abdominis segmentum primun
supra rufum, antice nigro-emarginatum, postice flavo-mar-
ginatum. ‘Tibiz tarsique rufescentes.
' Q ac 6 fere similiter colorata, sed clypeus ex magna parte
est niger, lateribus flavis aut aurantiacis, tibiis supra sepe longe
flavescentibus.
3 clypeus distincte nec profunde emarginatus, parce pune-
tatus, puncturatione minutissima, inter puncta majora, distincta.
2 clypeus plerumque levius, quam maris, emarginatus, et
crebrius, distinctissime punctatus. Mesonotum fortiter denseque
punctatum, postscutello inermi, propodeo rotundato, distincte
punctato, angulis pronotalibus parum distinctis. Abdominis
segmentum secundum dorsale pilis erectis, haud densis (et facile
detritis), ubique vestitum, basim versus (precipue ¢) sat con-
vexum. Segmentum 2 ventrale fortissime post suleum trans-
versum elevatum, elevatione summa prominula, Segmentorum
sequentium apices sepe aurantiace, aut plus minus pallide.
Alarum pars costalis distinete infuscata, partibus ceteris leviter
infuscatis. ¢ antenne 1] articulate ,articulis 3 ultimis minutis,
? haud plus quam solito insvassate. Long. 8-9 min.
This species somewhat resembles the preceding P. despectus,
but is probably not very closely allied, as the female—this being
the only sex known of the other—has the elypeus much less short
and much less convex, and the punctures are much less close.
The second ventral abdominal segment is more strongly raised,
the top of the truncation in the middle being of more pointed
form, the second dorsal segment less abruptly and strongly raised
from the base ete., and the coloration of the insect is brighter.
It is excessively like P. picteti Sauss. in the other section of the
genus. The emargination of the clypeus is very shallow in some
females, rather deeper in others.
’ erane :
Hab. Queensland, Cains, 2 9,1 $, July; Kuranda (Dodd),
1 9; Mackay and Kuranda, several examples (Turner).
WASP-GENUS PARALASTOR, 591
35, PARALASTOR ROSEOTINCTUS, sp. nD.
@. Nigra; abdominis segmentum primum supra (se. parte
declivi excepta) fere totum pailide aurantiacum, antice nigro-
emarginatum ; segmentum secundum fascia lata apicali, lateraliter
feve ad medium attingente, ornatum. Clypeus maculis duabus
curvatis magnis lateralibus, nonnunquam ad basim una con-
junctis, notatus; macula interantennalis magna, postorbitalis
parva, Antennarum articulus primus antice, aut totus, aut
basim versus, aurantiacus aut rufescens. Macule pronoti
magne, ad medium ejus longitudinem attingentes, nonnunquanmr
una conjuncte (sc. haud interrupte), et fasciam formantes.
Mesopleura notata ; tegule mesonotum juxta nigricantes. Scu-
tellum, postseutellum et propodeum aut utrinque notata, aut
tota nigra. Ale hyaline, parte costali parum late infuscata.
Tibie tarsique rufescentes. Color picturaticnis est insolitus,
pallide roseo-tinctus. Clypeus subnitidus, fortiter punctatus,
puncturatione minutissima interstitiali sat distincta, apice leviter
emarginato. Frons capitis densissime (nec. profunde) rugose
punctata. Anguli pronotales distincti, nee acuti. Mesonotum
sat grosse rugoso-punctatum, tegulis in parte exteriori haud
evidenter punctulatis, postscutello inermi. Abdominis seg-
mentum primum haud evidenter transversim impressum, rugoso-
punctatum ; secundum dorsale fortissime in formam conicam
elevatum ; ventrale, post suleum suum, distincte (nec fortissime)
supra partem suam basalem elevatum, elevatione summa late
rotundata, haudquaquam subacute prominula. Long. 8°5 mm.
The variation in the colour of the hind parts of the thorax is
unusual, but I have no doubt the two examples are of the same
species. The clothing of the thorax is not at all dense, so that
the sculpture is very easily seen. The very strong and pointed
elevation of the second dorsal segment of the abdomen is like that
of P. tuberculatus Sauss. alone of all the species known to me.
Hab. Australia, Swan River, 1 9 ; W. Australia, 1 2.
36, PARALASTOR XEROPHILUS, sp. n.
dé. Niger; clypeus, macula interantennalis, lineaque orbitalis,
in oculorum sinus extensa, flavescentia, Antennarum articulus
primus antice aurantiacus, et plus minus flavescens. Macule
postoculares verticis minores, elongate, aurantiace. Pronotum,
desuper aspectum, fere totum lete aurantiacum, propodeo
utrinque macula parva (an semper ?) rufescente ornato, Femora,
tibie tarsique aurantiaca. Abdominis segmentum primum
dorsaliter totum aurantiacum, hoe colore in partem declivem
extenso, basi nigra; segmentum secundum late aurantiaco-
fasciatum, fascia fere recta, aut tantum antice paullo concavo ;
Segmenta 2 sequentia aurantiaco-fasciata. Clypeus distincte,
nee profunde, emarginatus, haud dense, grossius sed leviter,
punctatus, et inter puncta majora dense minutissime punctulatus,
argenteo-pubescens. Frons capitis, cum mesonoto et propodeo,
592 DR. RB. C. L. PERKINS ON THE
quam densissime rugoso-punctata, postscutello inermi. Anguli
pronotales distincti, truncationis margine distincte elevato.
Tegule in parte exteriore, qua sunt latissime, impunctate vel
fere impunctate, puncturatione minuta absenti. Ale sub-
infuscate, parte costali saturatiore, basim versus flavescente.
Abdominis seementum primum rugoso-punctatum, medium im-
pressum ; secundum dorsale basim versus subfortiter convexuin ;
ventrale, post suleum costatum, fortiter elevatum, elevatione
summa paullo prominula, ibique grosse rugoso-punctatum ; seg-
menta apicalia tomentosa, ultimo pilis paucis brevioribus sub-
erectis vestito, minutissime dense punctulato, haud impresso.
Long. cirea 11 mm.
Hab. Central Australia, Hermansburg (17. J. Hillier), 1 ¢o.
37. PARALASTOR PLACENS, sp. n.
2. Nigra, clypeo, macula interantennali, et duabus verticis
postocularibus, pronoto, maculis subtegularibus, duabus magnis
scutellaribus et propodealibus, duabus postscutellaribus, abdominis
segmentum primum, triangulo elongato nigro basali, cujus apex
superficiem superiorem intrat, excepto, margine postico secundi
sat late, femoribus, tibiis tarsisque aurantiacis aut rufo-
aurantiacis. Fascia segmenti secundi media antice excisa, seg-
mentis sequentibus nigris. Tegule ex majore parte aurantiace,
Ale subhyaline, costali parte infuscata. Clypeus nitidus, sub-
tiliter perparce punctatus, punctis minutissimis etiam pareis,
apice distincte, nec profunde, emarginato. Frons capitis grosse
punctata, punctis sat distinctis. Pronoti truncatio haud mar-
ginata. Mesonotum dense subrugosim distincte punctatum,
punctis scutellaribus remotioribus. Postscutellum distincte
punctatum, inerme. Tegule polite, puncturatione minutissima
in parte exteriore carentes. Abdominis segmentum primuwn
subirregulariter punctatum, medium impressum: secundum
dorsale sat elongatum, lateribus perpaullo rotundatis, fascia
apicali excepta, tomento nigerrima; ventrale, post sulcum,
fortissime abrupte elevatum, post elevationem haud grosse
punctatum. Long. cirea 9 mm.
Hab. Australia, Swan River, 1 9.
38. PARALASTOR ERIURGUS Sauss.
The only examples of this large and robust species that I have
seen with a definite locality-label are from Queensland. The
apex of the clypeus is very distinctly margined, at least in
the female.
The male has the usual 11-jointed antennee, with the three
apical joints minute. The third and following ventral segments
of the abdomen are quite densely hairy, the apical segment
bearing quite long erect hairs, being finely punctured and without
an impression. The clypeus is densely pubescent, the larger
punctures obsolescent, the minute puncturation dense. The
WASP-GENUS PARALASTOR. 593
tegule in both sexes have a close minute puncturation on their
outer portion. The amount of black on the clypeus of the female
varies and probably may be altogether absent, while in some
the orange colour is reduced to a wide curved or horseshoe-
shaped mark. ‘The single male has only a small black spot near
the apex.
Hab. Queensland, Brisbane (Hacker), 4 92 taken singly in
JanweetebreMarchy and. April, 9 Hou other 9 and 1 ¢
examined.
39. PARALASTOR ARGENTIFRONS Sunith.
This species structurally seems to be almost identical with
P. eriurgus and is perhaps only a southern race of the Queensland
form, but is very different in superficial appearance. It varies
greatly in the width of the abdominal fasciz of the first two
segments. Of the 13 examples before me I do not think that
there are even two that are alike in the pattern of coloration of
the second dorsal segment. One example, in which the black
forms an equilateral basal triangle, bears a MS. label, ‘“ saucius
Sauss. Type.” The clypeus in the single male bears a black median
apical spot, in the female it may be all black or marked at the
base ; it also varies in puncturation, and in some examples is quite
dull, in others more shining. ‘The pronotal spots may be absent.
In some examples the upper edge of the truncation of the second
ventral segment is less pointed in the middle (as in the type) than
in others, but I have examples taken in company which differ in
this respect, but not otherwise. The silvery pubescence of the
sides of the clypeus and adjoining orbits is more distinct and
conspicuous 1n some than in others. The male characters are as
in P. eriurgus.
Hab. Adelaide (my collection), 3 9, 19 & 1 3 (Oxford
Museum); 8. Australia, 1 @ (Oxford Mus.) and 1 @ (British
Museum); Victoria (/rench), 4 2; 2 9 without special locality.
40, PARALASTOR SUBOBSCURUS, Sp. n.
3. Niger, clypeo flavo, medio nigro-punctato, macula elongata
interantennali maculaque parva verticis postoculari, flavis.
Macule pronotales, mesopleurales, et scutellares aurantiace,
Tegule aurantiaco-marginate. Abdominis segmenta 2 basalia
postice rufo-marginata (vix aurantiaca) fasciis fere rectis, minus
latis. Tibiee tarsique ferruginei aut rufescentes. Ale infuscatee,
parte costali saturatiore.
Species villosa, P. eriwrgo et argentifronte angustior. Clypeus
fortiter emarginatus, nitidus, remote punctatus, minus pubescens.
Frons rugoso-punctata. Pronoti truncatio haud evidenter mar-
ginata, angulis tamen lateralibus distinctis. Mesonotum grosse
punctatum, punctis distinctis, minus confluentibus. Postscutellum
inerme. Propodeum distincte punctatum, Abdominis segmentum
primum subequaliter punctatum, pilis longis vestitum ; secundum
594 DR. R. C. L. PERKINS ON THE
dorsale pilis brevibus erectis sparsim vestitum, dense nigro-
tomentosum; ventrale fortissime post: suleum transversum
elevatum, elevatione summa acutius prominula, puncturatione
vix grossa aut densa; segmentum ultimum, cum precedentibus,
pilis sat longis erectis vestitum. ‘Tegule in parte exteriore
minutissime punctulate, punctis subobsoletis. Long. 9°5 mm.
Hab, North Queensland (Dodd), 1 3.
4]. PARALASTOR SIMPLEX, sp. n.
2. Nigra, macula interantennali, aliisque duabus minutis
postocularibus aurantiacis. Pronotum et mesopleura maculis
magnis aurantiacis ornata. Scutellum juxta marginem suuin
posteriorem et postscutellum fascia interrupta aurantiaca ornata,
Yeoule pallide; alee subflavo-infuscate. Tibi tarsique ferru-
ginei, illisaut externe aut subtus nigricantibus, articulis tarsorum
nonnullis etiam plus minusve infuseatis. Abdominis segmenta 2
basalia aurantiaco-fasciata, fasciis haud latis, fere zequalibus.
Clypeus levissime emarginatus, parce subtilins punctatus, inter
puncta majora parce subtilissime punctulatus, opacus. Pronoti
truncatio (parte media excepta) fortiter elevato- marginata.
Mesonotum fortiter densissime punctatum, postscutello inermi.
Abdominis segmentum primum quam densissine punctatum ;
secundum dorsale dense nigro-tomentosum, basi haud convexim
elevata; ventrale, post suleum transversum, fortissime elevatuin,
elevatione summa conspicue prominente. Tegule opace, punctis
paucis majoribus exceptis, impunctate. Long. 13 mm.
This appears to be a very distinct species, but the single
example is much abraded, and the flavescent appearance of the
wings may be partially due to age, but on the other hand these
may be still yellower in fresh examples, as the stigma itself is
more or less pale in colour.
Hab. Albany (Brewer), 1 9.
42. PARALASTOR MIMUS, Sp. n.
@. Angusta, nigra; macula interantennalis, cum duabus elon-
gatis postocularibus, pronotum, tegule, scutellum, postscutellum,
propodeum, macule mesopleurales, abdominis segmentum primum
(parte decliva majore excepta), femora, tibiz tarsique, nonnunquam
etiam coxe et trochanteres, rufa aut flavo-rufa. Clypeus aut
totus niger, aut totus rufescens, aut niger, rufo-tinctus. Ale,
costali parte excepta, parum infuscate. Clypeus distinete emar-
ginatus, punctis majoribus et minutissimis commixtis signatus,
pubescens. Mesonotum postice irregulariter punctatum ; post-
scutellum inerme. Tegule minutissime punctate, puncta com-
pliva majora nonnunquam ferentes. Abdominis segmentum
prmum breviter campanulatum, vix densissime punctatum ;
secundum ad basim conspicue angustatum, lateribus fere rectis
aut parum rotundatis, satis foneuu basim versus fortiter con-
vexum. Segmentum 2 ventrale post sulcum transversum
WASP-GENUS PARALASTOR. 695
fortissime elevatum, elevatione summa haudquaquam acute
prominente, fortiter punctatum. Long. circiter 9 mm.
Very distinct from any of the preceding species by its colour,
narrow elongate form, the longer first abdominal segment, lone
subparallel- -sided second segment, ete. It might almost be placed
in the other division of the! species here described, and is included
in the tables of both divisions.
Hab. Australia, Swan River, 3 2 (two in the Oxford and one
in the British Museum); 1 @2 in very bad condition is labelled
‘** New South Wales,” probably in error.
43. PARALASTOR IMiITATOR, sp.n. (PI. I. fig. 10.)
2. Preecedenti simillima, sed minor, tegulis grosse punctatis,
puncturatione minutissima absente, antennis brevibus et crassis
facile distinguenda.
Form, colour, and general appearance almost exactly as in
P. mimus. In the single specimen the clypeus is black on about
the apical third, its apex is very lightly emarginate. The fourth,
fifth, and sixth antennal joints are extremely strongly transverse.
The tegule are very shining, conspicuously, coarsely, and deeply
punctured, and without the numerous minute punctures that
form the chief sculpture in P?. mimus. The wings are still clearer,
with very little infuscation along the costa. ‘The basal abdominal
segment is rather shorter and the second less abruptly narrowed
at the base. The second ventral segment 1s considerably less
strongly raised behind the transverse sulcus. Long. 7°D mm.
This species is very distinct.
Hab. Australia, Champion Bay, | @.
44, PARALASTOR DEBILIS, sp. n. | (CEM, dU, dite, 20.)
3. Niger, clypeo, antennarum articulo primo antice, maculaque
postoculari minuta, flavescentibus. Pronoti fascia, utrinque
dilatata, pars tegularum exterior, note parve scutelli due, fascia
sat lata postscutellaris, abdominis segmentum primum, parte
declivi excepta, faseia secundi apicalis, aurantiaca aut aurantiaco-
rufa. Tibiee ferruginez, tarsis ex majore parte fusco-brunneis.
Clypeus brevis, sat convexus, apice levissime emarginato, dense
argenteo-pubescens. Frons capitis pilis crebre vestita. Pronoti
truncatio fere recta, aut perpaullo concava, distincte elevato-
marginata. Mesonotum densissime fortiter punctatum, scutello
similiter punctato, postscutello inermi. Tegule nitide, grosse
et conspicue punctate. Abdominis segmentum primum, quam
densissime, grosse punctatum, secundum etiam densissime ubigue
punctatum, basi fortiter convexim elevata. Segmentum 2
ventrale post suleum transversum distincte (sed haud fortissime)
supra partem basalem suam elevatum, elevatione summa haud-
quaquam acutius producta, post hance late et conspicue im-
pressum, et crebre punctatum; segmenta apicalia appresse
596 DR. R. C. L. PERKINS ON THE
pubescentia, ultimo pilis paucis longiovibus erectis vestito.
Long. circa 7 mm.
The pronotal band at its widest hardly reaches the middle of
the length of the pronotum; the second abdominal segment is
thinly clothed with more or less erect hairs, its apical fascia
is not dilated at the sides, and only moderately wide, not
occupying more than about one-fifth of the segment.
Hab. West Australia, Swan River, 1869, 1 ¢ (de Boulay), in
the Oxford Museum.
45, PARALASTOR EUGONIAS, sp. n.
3. Niger; clypeus maculaque interantennalis elongata auran-
tiaco-flavescentes. Pronoti macule due, et tegule ex majore
parte, rufescentes. Segmentum abdominis primum et secundum
postice rufo-marginata, fascia prima latera versus angustata,
secunda utrinque valde dilatata, ibique ante aut ad medium
segmenti longitudinem extensa, ceteris segmentis (saltem ex
majore parte) rufis aut aurantiacis. Tibiz tarsique rufescentes.
Species minor, sed haud gracilis. Clypeus lunulato-emarginatus,
ex majore parte subdeplanatus, distincte, remote, haud grosse
punctatus, basi lateribusque suis argenteo-pubescens. Frons
cum thorace crebre pilosa. Mesonotum dense punctatum,
scutello inermi. Tegule grosse et conspicue punctate. Ab-
dominis segmentum secundum dorsale dense, grossius punctatum,
parte basali fortiter convexa; ventrale, post sulcum, fortiter
abrupte elevatum, et postice grosse denseque punctatum., Anguli
posteriores segmenti 2 dorsalis conspicue acute producti. Long.
circiter 8 mm.
Hab. Adelaide (A. K. Davis), 1 6.
46. PARALASTOR MICROGONIAS, sp. n.
3. Preecedenti simillimus, clypeo breviore et angulis segmenti 2
dorsalis abdominis posterioribus perpaullo productis distinguendus,
Long. circiter 7 mm.
Exactly like the preceding in general appearance, but the
elypeus is distinctly less produced apically and less flattened, and
its puncturation rather coarser. The lateral angles of the hind
margin are only very slightly produced backwards. The wings
are hardly infuscate, except along the costa, where the infuscation
is dark and conspicuous. In the preceding species the wings are
folded and appear dark, but this may be only due to the folding.
Both species have a small postocular spot on each side of the head,
which was not mentioned in my description of the preceding,
In both, the apical antennal joints are excessively minute,
forming a sort of small tubercle in the concavity of the eighth, and
it is ditticult to tell whether two or three joints are here present,
but I have satisfied myself that there are three, so that the
antenne are normal for the genus, 11-jointed.
Hab. Adelaide, 1 ¢ (British Museum).
WASP-GENUS PARALASTOR, 597
47, PARALASTOR SANGUINEUS Sauss.
I examined the type of this species in the British Museum, but
unfortunately have not been able to make an actual comparison
between it and the two preceding, one or other of which may be
only a variety of sanguineus. It has the apex of the clypeus much
more widely black, in the preceding only the actual margin being
black. From a rough sketch that I made, it is also, possibly, more
produced. The red apical band of the second dorsal segment
of the abdomen is produced further towards the base on each side.
These colour-characters, however, would of themselves be in-
sufficient for specific distinction. I did not observe the hind
angles of the second dorsal segment.
It certainly appears probable that one or other of these males
belongs to the female described as vulneratus Sauss.. but the very
different clypeus forbids us to associate them without further
evidence. It is usual for the male clypeus to be more deeply
emarginate than that of the female, whereas in the males and
females in question, the clypeus of the female is extraordinarily
deeply emarginate, far more strongly than in the male. Never-
theless, I suspect that sanguinews Sauss. and vulneratus Sauss.
are sexes of one species.
Hab. Australia (British Museum).
48, PARALASTOR ARGYRIAS, Sp. n.
3 colore P. sanguineo Sauss., congruens, sed clypeo toto flavo,
antennarum articulo primo antice flavescente, maculisque pro-
notalibus magnis, fasciam vix interruptam, utrinque dilatatam,
formantibus. Ale hyaline, costali parte infuscata. Clypeus fere
zqualiter conspicue convexus, parum profunde et fere equaliter
punctatus, puncturatione interstitiali crebra, ubique argenteo-
pubescens. Oculorum sinus utrinque flavo-notatus. Anguli pro-
notales parum distincti. Mesonotum dense punctatum, scutello
inermi, propodeo brevi et rotundato, tegulis grossissime punctatis.
Abdominis Segmentum primum dense, distincte punctatum,
lateribus suis, cum lateribus basalis partis secundi, conspicue
argenteo-tomentosis. Segmentum secundum dorsale latera versus
copiose, sed haud densissime, punctatum, basali parte fortiter
supra segmentum primum convexim elevata, fascia apicali utrinque
fere ad medium segmentum attingente, antice emarginata; seg-
mentum 2 ventrale, post sulcum transversum, fortiter abrupte
elevatum, post elevationem summam depressum, puncturatione
distincta, haud densa; segmentum septimum pubescentia decum-
bente vestitum, pilis longis erectis carens. Long. circiter 9 mm.
A very distinct species structurally, but resembling P. ewgonias
in colour and pattern, except for the yellow front of the scape
and the much greater development of the pronotal markings.
Hab. New South Wales, Wagga (Froggatt), 1 3g.
598 DR. R. C. L. PERKINS ON THE
A9, PARALASTOR EUCLIDIAS, Sp. n.
d picturatione fere P. argyriw ornatus, sed fascia segmenti 2
abdominalis fere ad basim utrinque protracta. Clypeus, anten-
narum articelus primus antice, macula elongata iterantennalis,
due postoculares, flavescentia aut flavo-aurantiaca. Macule due
pronotales aurantiace. Tegule incomplete rufo-marginate. Ab-
dominis segmentum primum postice late aurantiaco-fasciatum,
fascia antice emarginata ; fascia secundi fere ad basim segmenti
utringue producta (sc. segmentum secundum anrantiacum, et
triangulo magno nigro basali ornatum). ‘Tibize, tarsi apicesque
femorum aurantiaco-fulvescentes. Segmentum abdominalis ter-
tium dorsale, cum sequentibus, aurantiacum, segmentis ventralibus
(que post secundum posita sunt) atris, apicibus pallidioribus.
Clypeus sat longus, distincte emarginatus, obscure sparsissime
punctatus, et ubique densissime et minutissime punctulatus,
ubique argenteo-pubescens. Frons capitis dense, nee profunde
punctata. Mesonotum crebre punctatum, scutello sat convexo,
postscutello inermi, tegulis grosse punctatis, propodeo utrinque
rotundato. Abdominis segmentum primum rugose punctatum ;
secundum (in aspectu laterali) basim versus fere equaliter
curvatum, sive convexum, latera versus punctis Inter se ex magna
parte distinctis, haud ubique rugosim confluentibus. Abdominis
segmentum 2 ventrale post suleum suum transversum, fortissime
elevatum, elevatione summa conspicue prominente, post eleva-
tionem depressum et subfortiter sed vix grosse punctatum ;
segmentum ultimum latum, brevissime pubescens. Ale, costali
parte excepta, levissime infuscate. Antenne 11-articulate,
Long. circiter 10 mm.
As in many species, there is a silvery pubescence, conspicuous
in some aspects, along the lower part of the inner orbits and
filling the sinus of the eyes.
Hab. Victoria, Gippsland (Froggatt).
50. PARALASTOR OCCIDENTALIS, sp. n.
3. Niger, clypeo, macula elongata interantennali, articulo primo
antennarum antice, duabusque notis postocularibus, aurantiacis.
Pronotum, angulis deflexis inferioribus exceptis, totum, scutellum
postscutellumque, macula magna mesopleuralis, coxe (basi ex-
cepta), femora, tibiz tarsisque, cum lateribus propodei, duabusque
maculis mesonotalibus triangularibus, rufescentibus, vix auran-
tiacis. Tegule partibus thoracis ceteris pallidius colorate.
Abdominis segmentum primum (parte declivi excepta), fasciaque,
minus lata, apicalis segmenti secundi, rufa. Clypeus sat longus,
apice distincte, leviter emarginatus, densissime munutissime
punctulatus, punctis majoribus paucis interspersis, argenteo-
tomentosus. rons capitis rugose punctata. Pronotum fere
recte truncatum, truncatione bene elevato-marginata. Pronotum,
mesonotum et scutellum densissime punctata, scutello haud evi-
denta armato. Tegule profunde et grosse punctate. Abdominis
WASP-GENUS PARALASTOR. 599
segmentum primum densissime rugoso-punctatum, secundum
basim versus haud fortiter convexim elevatum, a latere visum,
tantum mediocriter convexum. Segmentum 2 ventrale, post-
suleum, fortissime abrupte elevatum, elevatione summa conspicue
prominente, sive tuberculata; segmentum ultimum Jatum, sub-
depressum, brevissime pubescens. Alarum pars costalis usque ad
stigma pallidum conspicue flavescens, parte cetera plus minusve
violaceo-nitente. Antenne Il-articulate. Long. 12 mm.
Hab. Australia, Swan River; 1 ¢ in the British Museum.
51. PARALASTOR ORIENTALIS, sp. n. (PI. I. fig. 21.)
Niger, clypeo, macula interantennali, duabusque postocularibus,
antennarum ¢ articulo primo antice, pronoto supra usque ad
tegulas, scutelli et postscutelli aut fascia completa aut maculis
duabus, propodeo utrinque, abdominis segmento primo (parte
declivi except) fere toto, apicali fascia secundi cum sequentibus,
tibiis tarsisque et sepe parte femorum majori, aurantiacis.
Clypeus et macula interantennalis sape, quam macule cetere,
flavescentiores. Ale distincte ubique infuscatee, costali parte
saturatiore. Clypeus distincte, nec profunde emarginatus,
fortiter, ¢ multo obscurius, punctatus, puncturatione minutissima
densa et conspicua. Mesonotum dense rugoso-punctatum, scutello
inermi, tegulis fortiter punctatis. Abdominis segmentum primum
distincte punctatum, pilis erectis perparce vestitum : ; segmentum
secundum tomento pernigrum, pilis erectis carens, fascia apical
latera versus paullo latiove. Segmentum 2 ventr ale, post sulcum,
fortissime elevatum, elevatione summa prominula, anterius
producta, segmento ultimo ¢ subdepressum, pubescens, pilis
erectis parum vestito. Long. 9-11 mm.
The orange colour varies in depth, being redder in some and
yellower in others. Sometimes there are two orange spots on the
mesonotum in front. There is also variation in the amount of
puncturation of the tegule. I have seen only six examples, and
four of these are without special locality.
Hab. New South Wales, 1 2 (British Museum); 1 2, Queens-
land, Bundaberg, taken by myself: 2 ¢ 2 2, Oxford Museum,
and 1 2, British Museum, without special locality.
52. PARALASTOR LACHESIS Sauss.
This species has the same coloration as P. orientalis, and may be
identical with it. Owing to the fact that the sculpture of the
tegulz is not referred to in Saussure’s description, I cannot place
it in my tables. The clypeus is said to be “ strongly emarginate,”
whereas the term is not as a rule used by Saussure for species
which have the clypeus no more strongly emarginate than
P. orientalis.
Hab. Yasmania, according to Saussure, but I have seen no
Tasmanian species with this style of ccloration, which is so
600 DR. R. CG. L. PERKINS ON THE
common on the mainland. Two other species, P. picteti and
P. tasmaniensis, recorded from Tasmania, are known to me only
from Queensland, and the possibility of error in Saussure’s
localities seems to be considerable.
53. PARALASTOR NAUTARUM Sauss.
This species, described on a single male, should probably be
placed near P. orientalis, as I have a note that the tegule are
very coarsely punctured, the clypeus distinctly, but not deeply,
emarginate. Superficially at least it is quite distinct from this
or allied forms, by the first segment of the abdomen being orange
with a black spot on its declivous basal portion, the second and
following ones being entirely black. The ventral surface of the
abdomen has a short dense pubescence.
Hab, Australia (British Museum),
54, PARALASTOR SOLITARIUS, sp.n. (PI. I. fig. 6.)
3g. P. orientali colore et vestitu simillimus, sed minor, orbitis
interioribus aurantiaco-fasciatis, maculis postocularibus magnis,
elongatis, ante medium oculorum marginem exteriorem extensis,
fascia pronotali haud ad tegulas extensa, tegulis ipsis inter puncta
majora minutissime punctulatis, abdominis segmento 2 ventrali,
post suleum, multo minus fortiter elevato, elevatione summa haud
aut vix antice producta, facillime distinguendus. Long. circiter
7 mm.
T have seen only one example of this species, which in general
appearance and sculpture resembles P. orientalis. The orange of
the basal segment is emarginate with black in the middle in front,
as in the other, but the second segment is less conspicuously deep
black, owing to the less dense tomentum, and the puncturation
therefore appears much more distinct. The tegule bear less
numerous punctures than is usual in this group.
Hab. Queensland, Bundaberg, 1 ¢, taken by myself.
55. PARALASTOR HILARIS, sp. n. (Pl. I. fig. 18.)
3. Caput flavum, colore flavo postoculari ab colore flavo fron-
tali separato, vertice nigro, hoc colore late in frontem producto.
Antennarum articulus primus, cum maxima mandibularum parte,
flavus. Pronotum, scutellum, postscutellum, propodeum utrin-
que, mesopleura antice, et tegule, flava, plus minusve hie illic
aurantiaco-tincta. Coxe, trochanteres, femora, tibie tarsique
flava, nonnullis in partibus brunnescentia. Abdominis seg-
mentum primum, parte declivi excepta, flavum, antice nigro- aut
brunneo-nigro emarginatum; segmentum secundum ad basim
brunneum, medium nigrum, postice sat late flavo-marginatum,
segmentis sequentibus plus minusve flavis. Antennarum flagellum
- subtus plus minusve rufescens. Clypeus sat longus, apice dis-
tincte emarginato, subeequaliter nec dense punctatus, puncturatione
WASP-GENUS PARALASTOR. 601
minutissina inter puncta majora densa. Frons capitis cum
thorace brevissime pilosus. Mesonotum nitidum, densissime,
‘ortiter punctatum, scutello inermi, tegulis grosse et profunde
vunctatis. Abdominis segmentum primum latissimum, fortiter
punctatum, pubescentia minutissima decumbente excepta, fere
nudum; segmentum secundum fortiter convexim elevatum, pilis
erectis carens, distincte crebreque punctatum. Segmentum 2
ventrale post sulcum sat abrupte, sed minus fortiter, supra partem
suam basalem elevatum, elevatione summa quasi incrassationem
transversam formante, post hane depressum et subtilius nec dense
punctatum ; segmentum ultimum pilis erectis evidenter vestitum,
leviter convexum, et subtiliter, nec dense, punctatum. Ale fere
hyaline, costali parte conspicue infuseata. Long. circiter 7 mm.
Hab. Northern Australia, Port Darwin; North Queensland ?
(Dodd). ,
56. PARALASTOR MACULIVENTRIS Sauss.
I think it probable that this species may be identical with my
P. elegans, described above. Saussure says nothing of the strongly
raised elongate clypeal carine, which Mr. Meade- Waldo informs
me occur in Saussure’s ty pe, as in P. elegans. He also informs
me that the type specimen is deformed, the sides of the face being
asymmetrical. The only difference, so far as I can judge, between
the two forms is that in P. maculiventris the apical band of the
first abdominal segment is entire, not disappearing before it
attains the lateral angles, that the head is said to be covered with
a blackish tomentum (pale in P. elegans, and conspicuously golden
in front), the ‘inner borders of the eyes are yellow” (only a
yellow spot in the sinus in P. elegans), while the latter has the
postscutellum hardly perceptibly tuberculate.
Hab. Australia. Type in the British Museum.
57. PARALASTOR BICARINATUS, Sp. N.
Niger, macula interantennali, duabus postocularibus, duabus
pronotalibus, maculis duabus, aut una magna, scutellaribus,
duabus propodealibus, tegulis, segmentum primum abdominis
(parte declivi excepta) ex majore parte, secundique fascia apical,
plus minusve antice utrinque producta, cum parte apicali seg-
mentorum sequentium, flavis. ¢ clypeus totus, antennarumque
articulus primus antice, flavi; 2 clypeus medius, cum lateribus
extra carinas niger. Al fere hyaline, costali parte infuscata,
stigmate atro-fusco. Tibi tarsique apicesque femorum flaves-
centes. Color flavus plus minus aurantiaco-tinctus. Clypeus
fortiter emarginatus et acute dentatus, parce punctatus, inter
puncta majora minutissime punctulatus, fortiter productus, et
duabus carinis elongatis, acute elevatis, munitus. Frons capitis
grosse et distincte punctata, interstitiis inter puncta minute
punctulatis. Mesonotum grosse punctatum, pilis erectis longi-
oribus vestitum, pronoto antice parum distincte marginato,
602 ‘DR. R, C. L. PERKINS ON THE
sentello inermi. Tegule parum nitide, minutissime obsoletim
punctulate, punctis nonnunquam tribus aut duobus majoribus
notandis. Abdominis segmentum primum minus for titer punc-
tatum ; secundum ad basin subfortiter convexum. Segmentum
2 ventrale post sulecum fortissime elevatum, elevatione summa sat
fortiter prominente; segmentum ultimum subconvexum, haud-
quaquam depressum, breviter pilosum. Long. 9-11 mm.
In the male the yellow of the first segment is nearly straight in
front, in the female it is emarginate in the middle. The ‘apical
band ah the second segment in the male does not reach the middle
of the segment, in the female it extends far beyond the middle
basally. “T think this is mere individual variation and probably
not a constant sexual difference.
Hab. Queensland, Mackay (Z'urner); 1 g and 1 Q.
58. PARALASTOR ATRIPENNIS, sp. n. (PI. I. fig. 13.)
2 color colori varietatum P. argentifrontis nounullarum per-
fecte similis, sed alis atris ubique violaceo-nitentibus primo
aspectu distinguenda. Macule parve interantennales cum
duabus postocularibus aurantiace. Segmentum primum ab-
dominis fascia apicali fere recta, haud dimidiam partem suam in
aspectu dorsali occupante, seeds fascia lata rufescente, antice
emarginata, latera versus ante medium segmentum protracta,
ornatum, exteris segmentis his fasciis colore fere similibus,
Tibie tarsique r -sufescentes. Tegulee obscurius rufescentes. Clypeus
fortiter emarginatus, dentibus lateralibus obtusis, inter hos
margine nonnullis in aspectibus perpaullo prominente, ut quasi
obscure tridentatus appareat, nitidus, antice fortius, postice
subtilius punctatus, et inter puncta majora minutissime punctu-
latus. Frons eapitis rugosa, punctis obscurissimis. Mesonotum
subopacum, puncturatione inwquali, punctis nonnullis grossioribus,
nonnullis minus grossis commixtis. Postseutellum medium paula
productum, sive ance ure tuberculatum. Tegule dense minutis-
sime punctate, punctis perpaucis majoribus presentibus. Ab-
dominis segmentum secundum ventrale post suleum transversum
fortissime abrupte elevatum, elevatione summa prominula, Long.
circa 14 mm.
IT have seen only one old example in dirty condition. It is
very distinct from any other that has similar coloration and
markings. The clypeus is more or less flat over most of its
surface, excepting the base and the decurved sides.
Hab. Adelaide (Wilson); 1 2 in the Oxford Museum.
59. PARALASTOR AUREOCINCTUS Guer.
There is a single female of this species in the British Museum,
bearing a label ce Alastor eriurgus, var.” Practically it resembles
eriurgus in colour, but in structure it is extremely close to the
preceding, to which it has no superficial hkeness. The tegule
WASP-GENUS PARALASTOR. 603
have the same dense, distinct, minute puncturation and the clypeus
is nearly of the same form, as also is the general sculpture. The
postscutellum has even less evidence of a tubercle than that of the
preceding species. Length about 15 mm.
Hab. Australia, without definite locality; 1 2 in the British
Museum. Judging from the coloration, one would expect to find
this species in Queensland.
60. PARALASTOR DENTIGER, sp. n. (PI. I. fig. 11.)
3 colore et picturatione fere P. mimi (q.v.) ornatus. Notex
postoculares minores, clypeo aurantiaco, margine apicali et non-
nunquam linea mediana nigris. Alee ubique infuscate. Clypeus
fortiter emarginatus, nitidus, parte media nitida, glabra, de-
planata, basi lateribusque decurvatis et dense argenteo-pubescenti-
bus. Frons dense et rugose punctata. Mesonotum densissime
punctatum, postscutello tuberculo spinoso conspicue armato,
tegulis puncturatione densissima et minutissima carentibus, sed
punctis nonnullis majoribus plerumque distinctis. Abdominis
segmentum primum transversim subdepressum, plus minusve
rugoso-punctatum, colore rufo antice profunde, sed anguste, nigro-
emarginato; segmentum secundum dorsale basim versus leviter
convexum ; ventrale, post sulcum suum transversum, fortissime
elevatum, elevatione summa prominente; segmentum ultimum
cum precedentibus pilis erectis evidenter vestitum. Antennz
1l-articulate, tribus apicalibus articulis minutis. Long. 11-5 mm.
This is a more robust species than P. mimus and imitator, which
it so much resembles superficially. The posterior face of the
propodeum is more extensively black than in the former. Struc-
turally it is entirely distinct from both.
Hab. Champion Bay and Swan River; 2 ¢ in the British
Museum.
61. PARALASTOR INFIMUS, Sp. n.
3 picturatione, forma, et colore P. eriurgo simillimus, sed apice
clypei haud distinctissime elevato-marginato, et scutello sat
conspicue tuberculato bene distinguendus. Clypeus leviter
emarginatus, grosse punctatus, nitidus, basi lateribusque dense
argenteo-pubescentibus. Frons dense rugoso-punctata. Tegulee
densissime et minutissime punctulate. Thorax sat grosse punc-
tatus, pilis erectis ubique conspicue vestitus. Abdominis seg-
mentum primum medium impressum, margine apicali excepto,
crebre punctatum. Abdominis segmentum 2 ventrale, post
suleum fortissime elevatum, subopacum, crebre punctatum,
elevatione summa prominula, hoe segmento cum sequentibus
pilis erectis fere xquilongis vestito, segmento ultimo spatium
medianum nitidius, lineis elevatulis definitum, prabente. Long.
12°5 mm.
Hab. Queensland, Brisbane (Hacker), 1 3d in December.
Proc. Zoou. Soc.—1914, No. XLII. 42
604 DR. R. C. L. PERKINS ON THE
62. PARALASTOR SUMMUS, Sp. N.
@.. Nigra, clypei lateribus deflexis (parte apicali excepta), macula
interantennali, duabus postocularibus, pronotalibus, mesopleura-
libus et scutellaribus cum parte tegularum, flavescentibus, aut
plus minus aurantiaco-flavis. Abdominis segmentum primum et
secundum late aurantiaco-marginata, fascia secunda utrinque
ante medium segmentum protracta, segmentis sequentibus, plus
minusve aurantiacis. Tibie tarsique rufescentes. Clypeus
(lateribus deflexis exceptis) deplanatus, nitidus, fortiter in parte
apicali punctatus, apice subfortiter emarginato et producto.
Frons cum thorace fusco-pilosa. Mesonotum grosse punctatum,
inter puncta minutissime punctulatum, sive quasi granulatum,
postscutello tuberculato, tegulis densissime punctulatis, opacis.
Ale subhyaline, parte costali infuscata. Abdominis segmentum
primum ex majore parte subtilius nec dense punctatum, secundum
basim versus minus fortiter convexum, tomento pernigro vestitum,
pilis erectis parcissimis. Segmentum 2 ventrale post suleum
suum fortissime elevatum, elevatione summa, desuper visa, sub-
angulata. Long. cireiter 12 mm.
This is a less robust species than the preceding, and the thorax
is notably longer. The flat portion of the clypeus becomes very
narrow posteriorly, being closely adapted to, and continuing the
plane of, the interantennal plate, when viewed from the side.
5 {oapn]
Hab. Queensland, Cairns, 1 °.
63. PARALASTOR MEDIUS, Sp. n.
2 precedenti cognatissima, clypeo breviore, hasim versus fortiter
sed sparsim punctato, propodeo utrinque maculato, distinguenda.
Very similar to the preceding, but the coloration is still
yellower (less orange), the clypeus is reddish apically and the
yellow colour of the sides extends to the apical teeth. The
yellow pronotal markings nearly reach the tegule, instead of
being confined to the front half; the band of the basal abdominal
segment is different in form, being emarginate in the middle of its
front margin, and the second is only a little widened at the sides ;
the legs are much more pale in colour. Strueturally, the clypeus
easily separates the two forms.
Hab. Queensland, Mackay, 1 2 (Zurner).
64. PARALASTOR EUSTOMUS, Sp. Nn.
QO. Nigra, clypeo utrinque late, macula elongata interantennali,
parte articuli primi antennalis antica, notis postorbitalibus duabus,
pronoto fere toto, tegulis, fascia scutellari et postscutellari, pro-
podeo utringue, femoribus, tibus tarsisque, segmento (parte
magna basali declivi excepta) primo, fasciaque apicali, vix dilatata,
secundi, cum segmentis sequentibus, aurantiaco-rufis. Clypeus
profundissime emarginatus, emarginatione, quam semicireulus,
profundiore, post hane depressus, nitidus, punctatus, lateribus
WASP-GENUS PARALASTOR. 605
deflexis opacis et pubescentibus. Frons capitis et thorax densius
pilosi, pilis minus longis. Pronotum antice fere recte truncatum,
parum distincte marginatum. Tegule dense minutissime punc-
tulate, fere opace. Postscutellum medium tuberculo spiniformi
armatum. Abdominis segmentum primum transversim subde-
pressum, haud grosse punctatum ; secundum dorsale, basim versus,
haud convexim elevatum ; ventrale, post sulcum transversum,
fortissime elevatum, fortiter nec dense punctatum. Ale in-
fuscatee. Long. circiter 12 mm.
Owing to the depression of the clypeus anteriorly, behind the
emar zination, two blunt or rounded carine are formed where the
sides are deflexed.
Hab. Australia, no special locality. Two old and dirty examples
in the British Museum.
65. PARALASTOR PSEUDOCHROMUS, sp. n. (PI. I. fig. 14.)
2 colore et picturatione P. argentifrontis Sm. ornata, et ut
probabile, eodem modo variabilis; structura a P. ewstomo vix
distinguenda. Long. 10°3-12 mm.
Two long curved spots on the sides of the clypeus and the
medio-frontal and postocular spots orange, those on the clypeus
reduced in size and redder in one example. Pronotal spots, which
are small, and the pale part of the tegule orange or red. Both
the first and second fascize of the abdomen are broad and laterally
dilated, the second at its middle reaching about to the middle of
the segment, and towards the sides considerably nearer still to
the base. The coloration is not the same as that of P. eustomes,
but is rather that described by Saussure as “ sanguineus.” Ex-
cepting that it appears to be of rather more slender and elongate
form, the species in structure is much like P. eustomus. Since
the two exawples, that I have seen, exhibit some variation, further
material is necessary to decide whether the present species and
P. eustomus are more than local colour-forms of one species.
Hab. Victoria (French); two examples.
66. PARALASTOR VULNERATUS Sauss.
This small but robust species appears to vary in the pattern of
the second abdominal segment, like P. argentifrons Sm., or rather,
I should say, like the examples I refer to that species. The
clypeus may be all black, or may have small or large, orange or
red, basal markings.
The extraordinary prolongation and flattening of the clypeus,
with its very deep emargination, and the flattening of the plate
between the antenne to adapt it to the base of the clypeus, sepa-
rate it from any other, except the following. The extremely deep
and coarse puncturation of the tegule, the form of the insect and
its sculpture, and the condition of the hind angles of the second
abdominal segment, are altogether like P. sanguineus Sauss.,
42%
606 DR. R. C. L. PERKINS ON THE
of which I suspect it may be the female. No two of the three
females are alike in detail, but the material is quite insufficient
for a proper understanding of the species. The clypeus is
normally extraordinarily dull, and I suspect this is due to some
special excretion, as a specimen I possess shows some more shining
spaces and some dull raised lumps, as if the excretion had not
been distributed in the usual even manner. ‘The scutellum is
unarmed, and the second ventral segment is very strongly and
abruptly raised behind the transverse suleature.
Hab. Victoria, 1 Q (French); 1 2, without special locality, in
the Oxford Museum; 1 9, Adelaide, in my collection.
67. PARALASTOR FALLAX, sp. 0.
2 picturatione et colore P. mimi (q.v.) et dentigert ornata.
Clypei pars deplanata antice nigra, postice rufa. Ale fere
hyaline, parte costali perconspicue infuscata. P. vulnerato
structura evidenter cognatissima, sed: major, clypeo nitidissimo,
mesonoto magis nitido, abdominis segmento 2 dorsali, basim
versus, leviter convexo, facile distinguenda. Long. 10 mm.
The markings are red, as in various other species from Western
Australia, which it entirely resembles superficially. I cannot be
certain whether the clypeus is always highly polished, as in the
single example described, or whether it may not be sometimes
dull, as in vulneratus, but in any case the species is easily
distinguished structurally by the small convexity of the second
abdominal segment, which in vulneratus, viewed laterally, rises
convexly above the first to a distinctly more conspicuous degree.
Hab. W. Australia, 1 2 (British Museum).
68. PARALASTOR BRISBANENSIS, Sp. N.
@. Nigra, clypeo utrinque, macula parva interantennali dua-
busque postocularibus, pronoto (desuper aspecto) ex majore parte,
tegulis, fascia lata scutellari, fascia postscutellari, macula utrinque
mesopleurali, abdominis segmento primo (parte declivi excepta)
ex majore parte, fascia secundi segmenti minus lata, fere recta,
cum marginibus posterioribus segmentorum sequentium, auran-
tiacis. Al infuscate, parte costali saturatiore. Clypeus leviter
late emarginatus, opacus, equaliter, subtiliter punctatus, et
lateribus deflexis argenteo-pubescentibus exceptis, ubique de-
planatus. Frons inter antennas deplanata, clypeo quam _perfec-
tissime adaptata, spatio depresso, ocellum anteriorem includente,
lineis curvatis levibus et elevatis late marginato. Mesonotum
grosse punctatum, postice tri-sulcatum, sulcis lateralibus ad
pronoti marginem interlorem extensis, postscutello inermi,
tegulis nitidis, sat conspicue grosse punctatis. Abdominis seg-
mentum primum profunde, sed minus grosse, punctatum, inter
puncta quam densissime punctulatum, sive granulatum. Seg-
mentum 2 dorsale pernigrum, densissime inter puncta minutissime
sculpturatum, basim versus fortiter convexum ; segmentum
WASP-GENUS PARALASTOR. 607
2 ventrale, post sulcum, fortissime elevatum, elevatione summa
prominente. Long. 11-5 mm.
The clypeus is wide and much less produced than in P. vul-
neratus, and is only feebly emarginate, but it is flattened in the
same perfect manner, and the “front, between the antenne, is.
similarly flattened. The raised smooth lines of the front of the
head are found only in this and the following allied form. The
femora all have a conspicuous yellow apical spot outwardly, which
contrasts strongly with the ferruginous tibiz and tarsi.
Hab. Queensland, Brisbane. I captured 1 @ on Christmas
Day, 1904.
69. PARALASTOR MACKAYENSIS, sp. n.
2 structura preecedenti fere similis, sed abdomine flavo-ornato,
pedibus pallidioribus, fascia prima abdominali antice sat profunde
nigro-emarginata, pronoti maculis multo minoribus, scutello post-
scutelloque binotatis, haud fasciatis, distinguenda.
In the very poor material at my disposal I see no structural
character that could be relied on, as constant, to separate this
form from the preceding, of which it is probably a more northern
race. In markings and in colour of the markings the two are
very distinct, the deep orange colour of brisbanensis being
replaced by a more ochreous or pale orange in mackayensis.
Hab. Queensland, Mackay, 2 2 (Turner); 1 2 from the same
locality from E. Saunders.
70. PARALASTOR PRINCEPS, Sp. n.
Niger, valde longipilosus, macula parva interantennali, dua-
busque minutis postocularibus, ornatus. Macule due ee
pronotales, fascizeque due abdominales, minus late, aurantiac
d clypeus cum parte articul antennarum primi antica Cee ne
Species grandis, pilis longis ubique conspicue vestita, capite
quadrato -incrassato, clypeo (precipue ¢) brevi, lato, apice
truncato, pilis longis erectis conspicue vestito; clypei 2 punctis
majoribus alisque minutis sat densis. Frons media dense
rugosim sculpturata, antennas versus et in sinu oculari utroque
pilis longis nigricantibus conspicue vestita. Mesonotum et
seutellum quam densissime sculpturata, postscutello inermi.
Abdominis segmentum primum pilis longis griseis, secundo
etiam et sequentibus pilis erectis longis et conspicuis vestitis.
Ale (costali parte excepta) exemplorum recentium parum
infuseate. Tegule subtiliter punctate, minutissime etiam
punctulate. Abdominis segmentum 2 ventrale post suleum
suum transversum fortissime elevatum, elevatione summa
prominente, pilis longissimis vestitum, segmento d ultimo
minutissime punctato, punctis paucis majoribus intermixtis,
pubescente, et pilis erectis sparsim vestito. Femora, tibie
tarsique nigra, aut nigricantia, tibiis posterioribus basin versus
608 DR. R. C. L. PERKINS ON THE
maculatis, pilis longis tenuibus sparsim vestitis. Long. 14-
15 mm.
This large hairy species with truncate clypeus cannot be
confused with any other. There is a good female in the British
Museum bearing a MS. label, ‘ Alastor australis,’ which is a
‘smaller species with ferruginous tibie and tarsi, and otherwise
different. There is a single male, deprived of the greater part
of its antenne, and a very old abraded female in the Oxford
Museum, which have the wings more infuscate or discoloured
by age.
Hab. W. Australia, 1 3, 2 2, as referred to above.
71. PARALASTOR SUBOLORIS, Sp. n.
Mas mari precedentis colore, vestitu et sculptura similis, sed
statura multo minore, thorace toto nigro, abdominis fascia prima
latissima, partem majorem segmenti, desuper aspecti, oecupante ;
tarsis omnibus tibiarumque apicibus testaceis, facillime distin-
guendus. ong. 10 mm.
Apart from the above characters, the apical ventral segment is
still broader, its minute puncturation is less dense, and the few
slightly larger punctures are hardly noticeable, while the second
ventral segment is much more finely punctured. A single
female in the British Museum may belong to this species. It
lacks the minute imterantennal spot of the male, but has two
small prothoracic ones. Its second dorsal segment is very long
and parallel-sided. It is in a dirty condition.
Hab. W. Australia, 1 ¢ in the Oxford Museum.
72. PARALASTOR OLORIS, sp. n. (PI. I. figs. 4 & 16.)
Mas mari P. suboloris forma, sculptura, vestitu et colore
simillimus, sed minor, antennarum articulo primo nigro, tibiis
tarsisque ferrugineis facile distinguendus.
A small narrow species, with the basal orange fascia deeply
emarginate in front, the 2nd ventral segment with large sparse
punctures, and the top of the truncation in the middle still more
strongly raised and prominent than in the preceding species.
The apical ventral segment is less wide, very dull from the
extreme density of the sculpture, which is hardly to be distin-
guished as puncturation, between the fine and sparse feeble
punctures, that can be distinguished as such. The clothing of
this segment consists of excessively short pubescence or tomentum,
long, erect hairs being absent. ‘he second ventral segment is
almost bare except just behind the tubercle, and the following
ones are merely tomentose. Antenne 1] -jointed, with the three
apical ones minute, as in P. suboloris.
Hab. Australia, Swan River; 1 ¢ in the British Museum.
73. PARALASTOR COMMUTATUS, sp. n.
ae ¢ :
©. Nigra, angustula, clypei macula basali magna, curvata,
macula interantennali, duabus postocularibus minutis, pronote
WASP-GENUS PARALASTOR. 609
ex parte maxima, maculis 2 rotundatis scutellaribus, duabus
minutis mesopleuralibus, fascia latissima segmenti 1 abdominalis,
faseia secundi fere recta et, quam fascia prima, multo minus lata,
tibiis tarsisque omnibus, rufescentibus. Clypeus truncatus, ex
majore parte dense, subtiliter ruguloso-punctatus. Mesonotum
posterius irregulariter aut vix dense punctatum, postscutello
inermi. Tegule vix nitide, ex majore parte subtilissime, vix
evidenter sculpturate, puncturatione minutissima distincta
carentes. Abdominis segmentum primum haud fortissime
transversum, parte declivi excepta fere totum rufum; secundum
elongatum, lateribus fere parallelis, basi haudquaquam convexim
elevata. Segmentum 2 ventrale post sulcum fortissime elevatum,
elevatione summa fortissime prominente, tuberculum fortem
formante. Long. 9 mm.
I have seen only a single female of this distinct species and
it is in dirty condition. The wings appear to be infuscate, but
this may be partly due to this condition. In any case the costal
portion is much darker than the rest.
Hab. Australia, Champion Bay; 1 9 in the British Museum.
74, PARALASTOR CARINATUS Sm. (? LATERITIUS Sauss., var.).
I have before me a single female from the Oxford Museum,
which I refer, as a slight variety, to this species. It has a pair of
small orange basal spots on the clypeus, and the black colour of the
second abdominal segment is more extensive, forming a basal
band, angulated in the middle and with irregular sinuations on
each side of this. The clypeus is truncate, punctate, and more or
less strigose, turned upwards on its apical portion, and with a
fovea in the middle adjoining the apical margin. There is in
some aspects a faint trace of tuberculation in the middle line of
the postscutellum. The second ventral segment is very strongly
raised behind the transverse sulcature, and somewhat pointedly
produced in the middle at the top of the truncation ; it is rather
finely and not very unevenly punctured, with very distinct
interstitial minute puncturation.
Hab, Adelaide, 1 9, Oxford Museum. The type is in the
British Museum and must nearly, if not altogether, agree with
the Oxford specimen, except in slight points of colour, which vary
shnilarly in other species.
7). PARALASTOR VIDUUS, Sp. n.
3. Niger, clypeo, articulo antennarum primo antice, maculis-
que 2 postocularibus, flavis. Pronotum colore aurantiaco-flavo
bimaculatum, scutello minute binotato. Abdominis segmenta
2 basalia sat late aurantiaco-fasciata, segmentis nonnullis sequen-
tibus etiam fasciatis. Clypeus nitidus, late truncatus, minus
inequaliter punctatus, pilis erectis, aut suberectis, pallidis
vestitus. Frons cum thorace pilis longis subfuscis conspicue
vestita. Pronoti truncatio evidenter marginata. ‘Tegulie nitidi,
610 DR. R. GC. L. PERKINS ON THE
ex maxima parte vix punctate. Postscutellum inerme. Abdominis
segmentum primum transversim depressum, margine posteriore
elevatulo, medium subfoveatum, puncturatione confusa ; secundum
dorsale pilis parcis longis erecte vestitum, fortissime convexim
elevatum ; ventrale, post suleum suum, fortiter elevatum, eleva-
tione summa haud prominente, cum sequentibus hirsutissimum,
segmento ultimo pilis erectis brevioribus sat dense vestito.
Antenne fortiter clavate, 12-articulate, articulis 4 ultimis parvis.
Long. circiter 11 mm.
Hab. Victoria, 1 $ (French). The specimen bears several
labels: “C.F. 8.00”; “Macedon 1.3.93”; ‘‘ Melbourne, Vic-
toria.”
76. PARALASTOR ODYNERIPENNIS, Sp. Nn.
2 colore fere precedentis, sed clypeo nigro, ad basim flavo-
aurantiaco-maculato, scutello et antennis nigris, tibiis nigri-
cantibus, tarsis ex majore parte atro-brunneis.
Structurally this appears to me to be almost identical with
P. carinatus, but the dark legs (the hind tibize inwardly at the
base being rufescent) easily distinguish it. The second cubital
cell is triangular and not at all petiolate. The second abdominal
fascia does not occupy half the length of the segment.
Hab. Victoria (French).
77. PARALASTOR ODYNEROIDES, sp. Nn.
3g colore P. vidui, sed macula interantennali elongata flava,
scutello nigro, maculis pronotalibus minutioribus aurantiacis,
distinguendus. Structura valde distinctus. Clypeus fere recte
truncatus, medius depressus, utrinque quasi rotundatim cari-
natus, obscure punctatus. Mesonotum grosse punctatum, post-
scutello inermi, tegulis nitentibus, ex majore parte obsoletissime
minutissime punctulatis. Abdominis segmentum primum fortiter
transversum, rugosissime grosse punctatum, parte declivi media
longitudinaliter carinata. Segmentum 2 dorsale fortiter basim
versus convexim elevatum, fascia utrinque ante medium seg-
mentum extensa, profunde rotundatim antice emarginata, seg-
mentis ceteris aurantiacis. Segmentum 2 ventrale post suleum
fortiter abrupte elevatum, elevatione summa subangulatim
producta; segmentum ultimum fortiter convexum, pilis erectis
vestitum. Antenne 11-articulate, articulis 3 ultimis minimis.
Cellula 2 cubitalis haud petiolata. Long, circiter 11 mm.
Hab. Australia; 1 ¢ in the Oxford Museum.
78. PARALASTOR MUTABILIS, sp.n. (PI. I. fig. 8.)
Q. Nigra, clypei macula basali transversa, antice emarginata,
duabus minutis postocularibus, duabus pronotalibus, tegularum
parte exteriori, fascia lata abdominalis segmenti primi, antice
emarginata, fascia secundi, aut fere simplici, aut ytrinque valde
WASP GENU PARALASTOR. 6i1
dilatata, et fere ad basum segmenti extensa, rufis aut rufescen-
tibus. Clypeus latus, truncatus, subeequaliter punctatus, et
minutissime punctulatus. Mesonotum crebre fortiter punc-
tatum, scutello inermi, tegulis nitidis, punctis majoribus non-
nullis, param fortiter impressis, presentibus. Ale infuscate,
parte costali saturatiore. Abdominis segmentum secundum
dorsale ad basim subfortiter convexum; ventrale, post suleum,
sat abrupte elevatum, parte antesulcali parti postsulcali alti-
tudine subquali, elevatione summa haudquaquam prominente.
Long. 9-10 mm.
With this species begins a series of forms with the hind part
of the second ventral segment raised only to about the height of
the basal portion, and with the clypeus always truncate apically.
Hab. Victoria (French); 2 2, Sept. 1901. The two examples
have a quite different pattern of the second abdominal segment,
and they are in a dirty condition.
79. PARALASTOR PLEBEIUS, sp. 1.
2. Nigra, macula clypeicurvata basali, notis 2 postocularibus,
duabus maculis pronotalibus, nonnunquam notis 2 scutellaribus,
margine postico segmentorum 2 basalium abdominis angustius,
flavis. Clypeus truncatus aut paullo antice rotundatus, fortiter
equaliter punctatus. Frons capitis cum mesonoto dense punc-
tata, pilisque brevioribus sat crebre vestita, scutello inermi.
Tegule nitide, parum distincte sculpturate. Abdominis seg-
mentum primum fortiter transversum; secundum ad_ basim
fortiter convexum, pilis erectis brevioribus ubique, nec dense,
vestitum ; secundum ventrale, ut in precedente formatum.
Al fusco-hyaline, parte costali multo magis infuscata. Tibi
tarsique rufescentes. Long. 9-10 mm.
Hab. Adelaide, 1 9, British Museum; 1 9, Oxford Museum ;
Victoria (Yrench), 1 ©.
80. PARALASTOR SUBPLEBEIUS, sp. n. (PI. I. fig. 15.)
2 precedenti cognatissima, sed major, fasciis abdominalibus
minus angustis, flavo-aurantiacis, segmento primo abdominis
conspicue transversim depresso, facile distinguenda. Long.
12 mm.
This is probably the representative in Queensland of P. plebeius,
but, apart from colour, the form of the basal abdominal segment
is too distinct to permit the two to be considered as varieties of
one species.
Hab. North Queensland (Dodd), 2 2.
81. PARALASTOR DEBILITATUS, Sp. n.
Color fere P. pusilli Sauss. et P. simulatoris mihi. Fascize
abdominales bicolores, antice rufescentes aut subaurantiacie,
postice flave. -clypeus flavus, 2 parte posteriori sola flava
612 DR. R. C. L. PERKINS ON THE
aut aurantiaca. Antennarum articulus primus ¢ antice flavus,
2 aut plus minus rufescens aut totus niger. Macule pronotales
bicolores. Scutellum seepius binotatum, tegulis subtestaceis.
‘Tibiz tarsique rufescentes. Species parva, angusta, clypeo et
segmento 2 ventrali ut in spp. precedentibus formatis, forma
gvacillimi facile distinguenda. Long. 7-8 mm.
In this and the three preceding species the antenne are very
clavate, the fifth joint being wider than long and the following
becoming still more transverse. The males, except that of
P. debilis, are not known. In this male the antenne are very
strongly clavate, the fifth joint not longer than its apical width,
the sixth and seventh strongly transverse. Inall there are twelve
joints, the three apical ones recurved to form a small hook, the
preceding one being larger, but small. The wings are nearly
hyaline, except along the costa, in one example, darker in the
others, which are in abraded condition. One example in the
Oxford Museum bears a MS. name, ‘‘ Alastor pusillus Sauss.,”
but this is clearly an error.
Hab. Victoria (French), 2 9, in the British Museum; 1 d,
Adelaide (Wilson), and 1 9 without locality, Oxford Museum.
82. PARALASTOR BRUNNEUS Sauss.
With this species is begun a short series of allied species, which
are quite distinct from any others by the form of the second ventral
segment, which, behind the longitudinally costate suleature, rises
very obliquely from this to a height not, or but little, exceeding
the level of the base of the segment. ‘he clypeus is always very
distinctly emarginate and plentifully and coarsely punctate, the
punctures in the male being, however, more feebly impressed than
in the female. The tegulz, over at least a considerable part of
their surface, always bear coarse punctures, the puncturation
being frequently rugose, but owing to the generally pale colour
of the surface, the puncturation is usually less conspicuous than
it would be were the tegule dark in colour.
Hab. Australia, 2 9 and the type in the British Museum
(unter).
83. PARALASTOR MULTICOLOR, Sjo ilge, (Uedle isatiegs S) 1G )5)
Precedenti cognatissimus, sed paullo minor; Q clypeo flavo
et macula verticis nigra, utrinque usque ad antennas producta,
facile distinguendus. Clypeus distincte dentato-emarginatus,
productus, minus latus, subgrosse, crebre punctatus, flavus.
Caput cum thorace perparce, vix evidenter, pilosum. Mesonotum
et scutellum densissime punctata, postscutello inermi. Abdominis
segmentum primum crebre rugosim punctatum, haud transversim
depressum ; secundum dorsale (a latere visum) fortiter sub-
equaliter convexim curvatum; ventrale, post suleum, oblique,
parum fortiter, elevatum, et late subdepressum; segmentum
ultimum ventrale ¢ brevissime pubescens. - Antenne 3, ut
WASP-GENUS PARALASTOR. 613
apparet, tantum 10-articulate, articulis 2 ultimis minutissimis.
Long. circiter 7-7-5 mm.
The colour of this little Ulee is variable. The colour of the
whole front of the head is pale (the clypeus always yellow, but
in other parts the yellow becomes partly or wholly orange), except
that in the female the black of the vertex is produced down
to the antenne in two broad stripes, which sometimes become
reddish towards the antenne. In the male the black colour is
prolonged downwards on each side for only a short distance,
being arcuately emarginate just in front of the anterior ocellus.
The posterior orbits are widely yellow or orange, and connected
with this same colour of the anterior ones. Around the ocelli
the vertex is black, but behind these may be either black or red.
The pronotum, senile, and propodeum (except in the posterior
concavity) are for the most part yellow or orange, the pronotum
sometimes more brownish or reddish posteriorly. The meso-
notum may be all black or nearly all red. ‘The second abdominal
segment is more or less brown or reddish in front of the orange-
yellow apical band, sometimes nearly entirely so. The basal
segment is yellow or orange-yellow apically, shading into brown
or reddish, black basally. The sides of the thorax may be black
or red, with yellow markings. Legs all pale, usually fulvous,
with bright yellow hnes on the tibiz and femora.
_ Very closely allied to P. brunneus, but the edges of the lateral
deflexed parts of the clypeus are not so sharply defined, and the
colour is very different.
T can Ce distinguish ten joints in the male antenne, and this
character may be peculiar to the group.
Hab. Port Darwin (Dodd).
84. PARALASTOR ANOSTREPTUS, Sp. ND.
@. Nigra, clypeo, antennarum articulis duabus basalibus,
orbitis anterioribus et posterioribus (haud interrupte), macula elon-
gata mediofrontal, a clypeo fere ad ocellum anteriorem extensa,
pronoto ex maxima parte, postscutello et scutello fere totis,
maculis magnis propodei lateralibus, parte magna mesopleurali,
fascia latissima segmenti abdominalis primi, fascia apicali,
utrinque subdilatata, secundi, cum pedibus omnibus, refescen-
tibus. Segmentum abdominis tertium et sequentia, plus minusve
colorata, Tegule brunnescentes, plus minus testacee. Clypeus
fortiter rugoso-punctatus, apice producto, parum lato, subfortiter
emarginatus, partibus deflexis argenteo-pubescentibus, utrinque
in parte apicali subcarinatus. Frons densissime 1ugoso-punctata,
pilis brevissimis vestita. Thorax haud aut vix pilosus, ubique
densissime punctatus, postscutello inermi, propodei lateribus
argenteo-tomentosis. Abdominis sesmentum primum cam panuli-
forme, sat longus, subdepressus, secundum densissime punc-
tatum, ex majore parte longitudinis fere equaliter et fortiter
convexim rotundatum, apicem suum versus fortiter transversim
614 DR. R. C. L. PERKINS ON THE
incrassatum, margine ipso valde abrupte dejecto. Ale ubique
fusco-nitid, parte costali saturatiore, basim versus subflavescente.
Long. 9 mm.
Hab. N.W. Australia, 8. Heywood I.; 1 2 in the British
Museum.
85. PARALASTOR DYSCRITIAS, Sp. n.
2 precedenti forma, colore et picturatione simillima, sed
abdominis segmento secundo ante incrassationem apicalem haud
transversim depresso, bene distinguenda. Long. 9 mm.
Differs slightly in colour from the preceding, as follows :—
The markings of the head and abdomen tend to a more orange-
red colour, and those behind the eyes do not extend beyond about
the middle of the length of these organs. There is a roundish
mesopleural spot beneath the tegule and only a very small one
beneath this; the tegule are much redder, the legs paler, more
fulvous. The apical dorsal segment is concolorous with the
orange ones preceding it. The third antennal joint is nearly
all red (only somewhat reddish in the other). These colour-
differences may be quite inconstant, but in dyscritias the tegule
are coarsely rugosely punctate over nearly the whole surface,
smooth, shining, and sparsely punctured over a considerable area
in anostreptus. The actual hind margin of the second segment is
more abruptly depressed in the latter, and its incrassation is
raised on the basal side from the general surface of the segment,
whereas in dyscritias it is not at all defined in front from the
rest of the surface.
Hab. Australia, 1 2 in the British Museum, without locality.
86. PARALASTOR XANTHOCHROMUS, sp. n.
@. Nigra, clypeo utrinque, macula interantennali, antennarum
articulo (ferrugineo) primo antice, maculis duabus elongatis
postocularibus, duabusque in sinu oculorum sitis, duabus pro-
notalibus, scutellaribus, mesopleuralibus et propodealibus flavis,
sive aurantiaco-flavis. Abdominis segmentum primum late
flavo-fasciatum, fascia profunde nigro-emarginata; secundum
cum sequentibus fascia apicali ornatum. Pedes fulvescentes,
partibus nonnullis flavescentibus. Long. 9 mm.
Extremely close to the preceding in structure, but entirely
different in appearance. The tegule have the same coarse
puncturation. The clypeus is a little more strongly produced,
and the front of the head rather more distinctly punctured,
while the apical margin of the second dorsal segment of the abdo-
men is less strongly and abruptly depressed.
A single male that I possess, from Townsville, does not agree
with either this or the preceding species, but is very like
dyscritias 2 in its markings, which, however, are orange-
coloured, not red, and there is a pair of large red spots towards
the base of the second ventral segment, not connected with the
WASP-GENUS PARALASTOR. 615
apical band. The structure of the second dorsal segment apically
is rather different from either species, but, in the absence of the
male sex of the others, it is too close to be worth describing as
distinct. This male was taken on March 15th, 1902, by Mr. F.
P. Dodd.
Hab. Queensiand, Cairns, 1 2.
87. PARALASTOR PICTETI Sauss.
With this species begins a series of allied forms, forming my
second division. It is Saussure’s division of Alastor picteti,
which he characterized as having the abdomen subpetiolate.
All the specimens of picteti that I have seen come from Queens-
land. The details of coloration in this species vary a little, and
I think none exactly agrees with Saussure’s description. He
gives “ Tasmania ” as the locality for his insect, but, as with
some other species already referred to, I doubt the correctness
of this. The tegule are for the most part smooth and polished.
Hab. Queensland, Bundaberg and Cairns, taken by myself ;
Mackay, three bad specimens (Z’wrner); Kuranda, May to June,
1913, eleven examples.
88. PARALASTOR CONSTRICTUS, Sp. n.
@. Nigra, clypei parte basali nigra, apice lateribusque flavis,
picturatione variabili. Macula interantennalis, margines oculares
antice posticeque plus minus—variabiles tamen sunt—pronoti
fascia irregularis anterior, ad tegulas lineatim utrinque producta,
macule scutellares, postscutellares, propodeales et mesopleurales,
segmentum abdominale (parte declivi excepta) ex majore parte
primum, fascia secundi lata cum duabus maculis magnis basalibus,
et segmentorum fascie sequentium, aurantiaca. Pedes fulves-
centes, femorum parte apicali sepe flavo-notata. Antennarum
articulus primus antice flavescens aut fulvus, flagello seepe subtus
rufescente. Clypeus truncatus aut vix emarginatus, margine
apicali recto aut perpaullo concavo, fortiter punctatus, punctura-
tione minuta interstitiali distinctissima. Frons cum thorace
pilis erectis brevioribus suberebre vestita, capite densissime punc-
tato. Mesonotum densissime grosse punctatum, puncturatione
distinctissima, sed subrugosa. Ale subhyaline, costali parte
fusco-flavescente, stigmate pallido, cellula marginali infumata.
Tegule ex magna parte leves, polite. Abdominis segmentum
primum grosse, sepe rugose punctatum, parte basali segmenti
secundi remote aut irregulariter punctata. Long. 9-11 mm.
I-have seen only five examples of this species and no two are
alike in details of markings or sculpture, but I think they
certainly all belong to a single species.
Hab. Queensland, Bundaberg, 2 9, taken by myself (June
1904); 1 2, Queensland (ea coll. H. Sawnders); Mackay (Turner),
2 Qe
In one of my Bundaberg examples the orbits are entirely
616 DR. R. GC. L. PERKINS ON THE
black, in the other they are yellow from the clypeus to the middle
of the ocular sinus; in Saunders’s example they nearly connect
with the postocular orbital stripes.
89. PARALASTOR MESOCHLORUS, sp. 0.
@ precedenti cognatissima, sed pronoti maculis haud ad
angulos humerales extensis, segmento 2 abdominalis ad basim
nigro, aut tantum maculis rufescentibus ornato, colore primi
leete aut pallide flavescente, distinguenda. Long. circiter 9 mm.
This species is again variable, and there are evidently two dis-
tinct forms of it, but the material is far too poor for a proper
understanding either of its definite specific characters or of its
variation. Of the typical form there are 2 males and | female.
In the female the clypeus is almost truncate, red at the apex
and along the sides, the antenne are red beneath, infuscate above,
the scape and following joint less so than the others. There is a
yellowish interantennal spot and a small orange one behind the
eyes. The pronotum is red-marked in front medially and this
colour extends back to the tegule very narrowly as a sutural line.
The scutellum, postscutellum, and propodeum have large red
markings, the tegule are reddish brown. Basal abdominal seg-
ment above bright (slightly orange) yellow. Second with a dull
orange band not occupying one-fourth of the segment, the
following ones also with apical b bands. Hind tibie dark at the
apex, at least inwardly. Mesopleura with a red spot.
Clypeus of the male lightly emarginate, yellow, becoming red-
dish basally at least in the middle, otherwise the head coloured
much like that of the female. Pronotum with red markings,
produced to the humeral angles and along the suture to the
tegule. The other thoracic markings are like those of the female,
iba bright yellow, a little orange- tinted, nearly concolorous with
the basal abdominal segment; second segment either all black
or coloured much as in the female, the basal spots reddish or
brownish, if present.
PARALASTOR MESOCHLORUS MESOCHLOROIDES, st. n.
Female clypeus as in the preceding, but the coloured parts are
yellow or partly yellow. Inner orbits with a yellow line from
the clypeus extending into the sinus. Pronotum only with two
obscure spots or faintly reddish on the suture medially. Thoracic
markings bright lemon-yellow, the black median part of the pro-
podeum less wide than in the male of the preceding, which these
females resemble in appearance. Basal abdominal segment nearly
concolorous with the thoracic markings. Second all black, the
apical margin at most obscurely alee Tegule bright yellow
inwardly, bad at the point of their deflexion less coarsely and
deeply punctured than in the ples cans form. Male not known.
Hab. Queensland, Mackay, 2 ¢, 1 2 typical form, in Sept. and
Oct. 1899 (Turner); race mesochloroides, 2 © at Kuranda, Feb.
and April, 1902 (Zurner).
WASP-GENUS PARALASTOR, 617
90. PARALASTOR DARWINIANUS, sp. n.
@. Nigra, clypeo cum parte mandibularum basali et anten-
narum articulo primo antice, macula mediana post clypeum, fere Ad
ocellum anteriorem extensa, orbitis interioribus et posterioribus
(haud una conjunctis), fascia transversa pronotali, ad tegulas
lineatim postice producta, utrinque dilatata, fere ad mesopleura
producta, macula mesopleurali, tegulis thoracem versus, lineis
duabus mesonotalibus elongatis, maculis 2 scutellaribus, post-
scutellaribus et propodealibus, margine postico sat late seemen-
torum abdominalium, flavis. Antenne cum parte majore pedum
brunnescentes. Abdominis segmentum primum et secundum
brunnea, plus minus atro-suffusa. Clypeus latus, minus fortiter
productus, nitidus, subtiliter punctatus, apice emarginato, margine
elevatulo. Fronscum thorace breviter erecte pilosa. Mesonotum
distincte, vix dense punctatum. Abdominis segmentum primum
haud ubique densissime punctatum, medium subfoveatum. Alze
subfusce, nitentes, stigmate pallido. Tegule leves, nitidissime.
Long. vix 10 mm.
A very distinct species. The yellow colour of the inner orbits
curves round on each side behind the ocelli, but the extremities
do not meet, and are well separated from the broad postocular
stripes.
Hab. Port Darwin, 1 2 (Dodd).
91. PARALASTOR COMPTUS, Sp. n.
3. Niger, clypeo, macula mediana excepta, mandibularum parte
basali, macula elongata frontali, orbitis interioribus ad sinus
oculorum et sinubus ipsis, macula parva verticali oculos juxta,
orbitis posterioribus lineatim ex majore parte, pronoti margine
antico, macula mesopleurali, tegulis ex magna parte, maculis
duabus scutellaribus, postscutellaribus et propodealibus, margine
postico segmentorum 2 basalium abdominis, tibize omnes externe,
et parte exteriore femorum intermediorum, flavis. Antennarum
flagellus subtus fulvescens, supra infuscatus, rufo-tinctus. Stigma
alarum atrum. Clypeus distinctissime angulatim emarginatus,
punctis majoribus sparsissimis, albo-pubescens. Mesonotum cum
scutello profunde grosse punctatum, punctis inter se distinc-
tissimis. Tegule ex majore parte leves et polite. Postscutelluin
cum propodeo distincte et profunde punctatum, punctis quam
mesonotalibus paullo minus grossis. Abdominis segmentum
primum grossissime punctatum. Segmentum 2 ventrale post
suleum late depressum, depressione antice elevato-marginata,
segmento ultimo pilis sparsis erectis longioribus vestito. Long.
7-8 mm.
_ Var. RUBESCENS, nov.
Pronotum ex majore parte et pars mesonoti media late, rufes-
centia. Fascia prima abdominalis vage rufo-marginata. Seg-
mentum secundum atro-brunneum.
618 DR. R. CG. L. PERKINS ON THE
I have seen only two males of this species, differing greatly in
colour, as above described. The antenne are 11-jointed, the
three apical ones very minute, and difficult to see in my speci-
mens, being sunk in the apical cavity of the eighth joint.
Hab. North Queensland, Herberton (Dodd).
92. PARALASTOR ALEXANDRIA, sp.n. (PI. I. figs. 7, 22.)
With this species begins a small series of forms, distinguished
at once from any of the preceding by the sculpture of the tegule,
which are either densely minutely punctured over nearly all their
surface or bear coarse and conspicuous punctures, with evident
fine ones between these, towards the outer margin on the anterior
part.
Brunneus, capitis vertice, nonnunquam etiam fronte, et rarius
parte mesonoti antica, nigris. Clypeus ¢ totus, 2 utrinque,
macula interantennali, orbitis internis et externis (nonnunquam
vix interrupte), maculis duabus pronotalibus, mesopleuralibus,
scutellaribus, postscutellaribus et propodealibus cum majore tegu-
larum parte, albo-flavis. Segmenta 2 basalia postice albido-flavo
marginata, fascia prima lata, segmentis sequentibus etiam plus
minusve fasciatis. Antenne subtus plus minus rufescentes,
articulo primo sepe antice albido-flavo, aut toto ferrugineo, aut
supra nigrescente, colore variabili. Ais hyaline, parte costali
plus minus infuseata, cellula marginali ex majore parte infuscata.
Tibize omnes cum femoribus anterioribus et intermediis albido-
lineatee. Clypeus distincte emarginatus, 2 remote punctatus,
interstitiis minutissime punctatis, d obscure punctatus, punctis
sat magnis, sed levissime impressis, ubique albo-pubescens. Caput
cum thorace tomentosus, hoc pilis erectis perpaucis aut nullis
vestito. _Mesonotum postice sparsim aut irregulariter punctatum,
superficie interstitiali conspicue minute punctata. Tegule crebre
et minutissime punctate. Abdominis segmenti secundi margo
apicalis fortiter depressus, latus; segmentum 2 ventrale antice
grosse, postice subtilius, punctatum. Long. 75-9 mm.
Hab. N. Australia, Alexandria, Dec. 1905,2 9, January, 1907,
1 Sg andl 9 (W. Stalker); Adelaide River,2 9: all in the British
Museum.
93. PARALASTOR ARENICOLA, Sp. n.
Q. Caput nigrum, clypeo rufescente, utrinque flavescente ;
lineis orbitalibus, in sinum oculorum a clypeo productis, maculaque
interantennali, flavescentibus, lineis postorbitalibus usque ad
verticem capitis extensis. Pronotum rufescens, antice albido- .
flavo aut aurantiaco-flavo colore variegatum ; mesonotum nigrum,
plus minusve rufo-suffusum aut rufo-notatum; scutellum, margine
antico excepto, cum postscutello rufescente, ambobus flavo-bino-
tatis. Propodeum rufescens, latera versus nonnunquam flavescens.
Mesopleura macula flavescente ornata, rufescentia aut nigro-
variegata. Pedes rufescentes, tibiis albido-flavo lineatis. Abdomen
WASP-GENUS PARALASTOR. 619
rufescens, segmento primo ante fasciam apicalem, latam, albido-
flavam, nonnunquam nigro, secundo fascia fere simplici lata ornato,
maculaque utrinque magna basali, pallide flava aut albida decorato.
Clypeus levissime, sed sat distincte,emarginatus. Long. 7°5 mm.
Allied to the preceding, with generally similar sculpture and
the apical margin of the second abdominal segment similarly de-
pressed and wide. Easily distinguished by the fairly numerous
and conspicuous large punctures of the tegule, and the rougher
postscutellum, which is rather differently formed,
Hab. Central Australia, Hermannsburg, 2 2 (4H. J. Hillier).
The specimens are not in very good condition, the one which is
not mutilated being probably somewhat discoloured.
94, PARALASTOR SIMILLIMUS, Sp. n.
Picturatio et color fere P. constricti (q. v.), sed statura minore.
Clypeus ¢ flavus, 2 macula parva nigrescente signatus, apice
truncato, aut vix evidenter emarginato.
2 clypeo obsoletius punctato (se. punctis parum fortiter im-
pressis), antennis crassioribus, tegulis opacis aut vix unitidis,
puncta grossiora ferentibus, distinguenda. Long. 8 mm.
As the female of this species is unique and the male of P. con-
strictus is unknown, I have only been able to compare this female
example with the latter species, and unfortunately I do not feel
quite sure that this single female belongs to the males that I a ssign
to simillimus. On account of the better material, I have taken a
male as the type and not this unique female. It lacks the pair of
large basal spots on the second abdominal segment at the base and
the tegule are less punctured than in the male. Seen in profile,
the second segment is much more evenly rounded above than in
P. constrictus, and this is also, to a less extent, true of the male,
The male has the tegule very coarsely punctured over nearly the
whole surface, the clypeal punctures are feebly impressed and the
apex of the clypeus is truncate or almost so. The general
sculpture is nearly the same as that of constrictus. The second
ventral segment is not flattened or slightly depressed behind the
transverse sulcus as it is in constrictus. The apical ventral
segment has a very short pubescence, which is decumbent or sub-
decumbent.
The species is quite distinct, and it is highly probable that the
sexes are correctly associated.
Hab. Queensland, Mackay (Zurner), 1 g, 1 9. I took the
former sex at Bundaberg, but no °.
95. PARALASTOR SYNCHROMUS, Sp. n.
Picturatio P. constricti picturationi fere similis, sed 2 clypeo
ruto, basim versus utrinque flavo-ornato, pronoto, desuper aspecto,
ex majore parte aut ubique colorato, rufo flavoque colore variegato,
scutello postscutelloque flavo-fasciatis (illo haud maculis bene
Proc. Zoou. Soc.—1914, No. XLIII. 43
620 DR. R. C. L. PERKINS ON THE
separatis ornato), fasciis integris, aut vix interruptis, maculis
propodealibus rufis aut flavo-rufis, segmento 2 abdominali maculis
basalibus carente, distinguenda. do ¢ tegulis minutissime evi-
denter punctulatis et clypeo distinctissime emarginato, facile
distinguendi ; a P. simillimo clypeo distincte emarginato, primo
aspectu separandi. Long. 8-9°5 mm.
The apical margin of the second abdominal segment is quite
strongly depressed and rather wide; ventrally behind the sulca-
ture it is depressed, and very remotely punctured. The apical
ventral segment of the male is subconvex and clothed with erect,
but not long, hairs.
Hab. Queensland, Mackay, ¢ 9, 6 examples (Zurner) ; Bunda-
berg, 1 ¢,1 @, and Cairns, 1 2, taken by myself.
96. PARALASTOR LEPTIAS, Sp. n.
Clypeus 3, macula interantennalis, linea orbitalis sinum
oculorum intrans (nonnunquam ex parte obliterata aut rufescens),
antennarum articulo primo ¢ antice, plerumque pallide flaves-
centia aut albo-flava. Pronotum ex magna, aut ex majore parte,
scutellum et postscutellum maculatim, propodeum latera versus,
rufescentia, hic illic flavo-variegata. Tegule plus minusve flayo-
notate. Abdominis segmenta 3 aut, 2 flavo-albido marginata,
fasciis his antice rufo-variegatis. Pedes colore variabiles, rufes-
centes plus minus flavo-albido-variegati, femoribus posterioribus
plerumque nigris. Mandibule rufescentes, ¢ albido-notate.
Clypeus 9, et antennarum articulus primus antice, rufescentes,
illo plerumque utrinque albido-maculato. Clypeus ¢ sepe plus
minus antice rufo-notatus. Mesopleura rufo-maculata, macula
nonnunquam plus minus flavescente. Ale, costali parte excepta,
parum infuscatee.—Species elongata, gracilis, abdominis segmento
primo campanuliformi, secundo elongato, angusto, lateribus sub-
parallelis, basim juxta angustatis. Clypeus subfortiter emar-
ginatus. Caput cum thorace pilis brevioribus erectis vestitum.
Mesonotum et scutellum grosse, distincte punctata, puncturatione
mesonotali postice irregulari aut minus densa. Abdominis seg-
mentum ventrale g apicale pilis brevissimis vestitum, leviter
convexum. Antenne ¢ 11-articulatze, articulis 3 ultimis minimis,
in conecavitate articuli 8 receptis. Long. 7°5-8°5 mm.
On account of its very narrow second segment of the abdomen,
this species appears rather out of place in this division, the first
segment appearing less small in comparison with the hinder
parts. It varies a good deal in details of coloration, the second
abdominal segment sometimes having two conspicuous rufescent
basal spots and sometimes lacking these. The lines along the
outer orbits are usually distinctly bicolorous, and are abbreviated
in some specimens. The tegule have a minute and plentiful
puncturation, and generally a few larger, but feeble, punctures
are noticeable.
Hab. Adelaide; collected for me by some of the late T. B,
Blackburn's pupils.
WASP-GENUS PARALASTOR. 621
97. PARALASTOR IGNOTUS, sp. n.
3. Clypeus, antennarum articulus primus antice, pronotum,
tegule, macula mesopleuralis, scutellum postscutellumque et pro-
podeum utrinque, aurantiaco-flavescentia. Abdominis segmentum
primum, plaga nigra basali excepta, cum pronoto fere concolor,
secundo fascia simili, irregulari, apicali, duabusque maculis
basalibus obscurioribus, ornato. Pedes ex majore parte aurantiaco-
flavescentes, femoribus posterioribus nigris. Stigma alarum
atrum. Antennarumarticulus 8 plus minus rufescens. Clypeus
subfortiter dentato-emarginatus, subcrebre punctatus, dense
argenteo-pubescens. Mesonotum fortiter punctatum, nitidum.
Abdominis segmentum ultimum ventrale pilis erectis, sat con-
spicuis, densius vestitum, subtilissime distincte punctatum.
Tegule nitide, puncturatione minuta distincta, paucis punctis
majoribus presentibus. Long. circiter 9 mm.
Hab. Australia, Swan River, 1 ¢.
98. PARALASTOR ICARIOIDES, sp.n. (Pl. I. fig. 2.)
Colore et picturatione fere P. mesochloro var. mesochloroidet
assimilis, maculis scutellaribus, propodealibus et postscutellaribus
cum abdominis segmento primo citrinis, sed tegulis minute et
copiose punctatis facillime distinguendus.
This species varies in details of coloration, the pronotum
above being sometimes nearly all red, sometimes only spotted
with red in front, and may have two quite distinct yellow spots
in the red colour. The clypeus may be entirely yellow in the male
or yellow only at the base, the rest being reddish or brownish
red. In the female it is usually red, sometimes, however, more or
less yellow at the sides towards the base. The inner orbits may
be yellow or red, or with the markings wanting. The second
abdominal segment is all black or may become piceous along the
apical margin, or with a brownish apical fascia. The wings are
vellowish along the costa, the marginal cell conspicuously infumate,
the stigma pale. The apical margin of the second abdominal seg-
ment is strongly depressed, the flattened margin wide. Antenne
brown, often paler beneath, sometimes nearly wholly fulvescent,
the scape yellow in front in the male. Length 7-9 mm.; the
former measurement from a small male.
I have examined fourteen examples. The species is quite
distinct from P. mesochlorus by the different scutellum, apart
from the tegular sculpture.
Hab. Queensland, Townsville (Dodd & Stalker); Kuranda near
Cairns (Zurner §& Dodd); Cairns, taken by myself.
Species que sedis incerte sunt, aut haud supra enumeratze.
I have in the first part (g. v.) of this paper suggested the
probable position of some of the following species or otherwise
referred to them :—
99. P. AUSTRALIS Sauss.
Hab. Australia.
43
622 DR. R. GC. L. PERKINS ON THE
100. P. GRUENTATUS Sauss.
Hab. Australia.
101. P. GR#FFEI.
Hab. Fiji; Ovalau.
102. P. smirut Sauss.
Hab. Australia.
103. P. HrRTIveNtTRIS Cameron.
Hab. New Guinea.
104. P. aBnormis* Bingham.
Hab. W. Australia.
PsEUDOZETHUS, gen. nov.
3. Clypeus transversus, subrotundatus, clypeo generis [schno-
celie fere similis. Mandibule breves, robustz, apice obliquo,
dentibus 4 armato, dentibus 2 intermediis minoribus. Palpi
maxillares 6-articulati, articulis 4 basalibus haud multo longi-
tudine inequalibus, his omnibus sat fortiter elongatis, articulo
quarto duobus ultimis, una conjunctis, vix breviore. Ligula
sat elongata, haud setosa, quam mentum circiter bis longior.
Palpi labiales 4-articulate, articulo primo gracili, perlongo, apice
dilatato, quam secundum (quod elongatum est, et sat robustum),
multo longiore ; articulis 2 ultimis elongatis, gracilibus, sub-
equalibus, una conjunctis secundo vix longioribus. Antenne
13-articulate, articulo ultimo uncum fortem formante. Pronotum
ad truncationem suam emarginatum, angulis distinctis, margine
acute elevato. Postseutellum postice truncatum, desuper visum
postice emarginatum, angulis lateralibus conspicue productis.
Propodeum medium impressum et profunde foveolatum, impres-
sione angustula, lateribus suis rotundatis. Abdomen ut in
nonnullis speciebus generis Zethi formatum (cf. 7. cwrulei-
pennis F.), Segmenti primi pars pedicellata sat elongata; pars
postpetiolaris subglobosa, aut valde supra convexa, ante marginem
apicalem profunde transversim sulcata. Segmentum secundum
longe campanulatum, basi valde constricta, quasi breviter pedi-
cellata.
Tibie intermediz duobus calcaribus distinctis instruct.
Alarum cellula 2 cubitalis supra brevissime petiolata. Hoe
excepto, neuratio fere /schnocelie. Tarsorum ungues bifide.
2 incognita.
* T have recently been able to examine this species. Bingham’s remarks (Tr. Ent.
Soc. 1912, p. 381) on the antennz are incorrect. In the examples he described, a
moderately strong le1is shows the antennz to be normal for Paralastor, i.e. 11-jointed.
The species should be placed next to P. orientalis.
WASP-GENUS PARALASTOR. 623
PsEUDOZETHUS AUSTRALENSIS, Sp. n.
Niger, clypeo, linea mandibulari, articulo antennarum primo,
pronoti fascia transversa, tegulis, maculis scutelli duabus, spinis
postscutellaribus, tibiis, tarsis, femorumque apicibus, fascia lata
postpetiolari et margine apicali segmentorum sequentium, colo-
ratis, partibus nonnullis colore flavo-aurantiaco, nonnullis rufo
decoratis. Thorax sat nitidus, grosse punctatus, propodeo rugo-
sim punctato et opaco. Abdominis pars postpetiolaris seomenti
primi fortiter punctatus, suleo apicali trifoveato. Segmentum
secundum fusco-tomentosum, medium juxta marginem apicalem
foveatum, subtilius remote punctatum, fascia apicali circa partem
quintam segmenti occupante. Ale flavo-hyaline, costam juxta
flavescentes. Long. 13 mm.
The clypeus and the mandibular lines are orange-yellow, the
other markings orange-red or red. The third abdominal segment
is less distinctly foveate at the middle apically than the preceding
one. Allied to Hlimus, but the spinose angles of the post-
scutellum and the Zethus-like form of the basal abdominal
segment are very distinctive.
Hab. N. Queensland, Australia (F. P. Dodd).
RHYNCHIUM AUSTRALENSE, Sp. n.
3. Aurantiaco-flavum, clypeo, et antennarum articulo primo
antice, flavis. Propodeum abdominisque segmenta 2 basalia
nigra aut nigricantia. Margo apicalis segmenti primi obscure,
secundi late, aurantiacus. Pedes posteriores ex majore parte
nigre. Ale flavescentes, apicibus late nigris. Apex clypei
levissime emarginatus, utrinque subcarinatus. Mesonotum minus
grosse punctatus, postice leve et impunctatum ; seutellum im-
punctatum ; ; propodeum rugosum, lateribus spinosis, concavitate
sua ex majore parte levi et nitida, postice transversim oblique
rugosa. Abdominis segmentum primum remote subtilius punc-
tatum, secundum similiter punctatum, fascia aurantiaca partem
circiter dimidiam apicalem occupante; segmentum apicale
ventrale, basi excepta, pilis longioribus erectis vestitum.
Long. 14mm. @ incognita.
In the single example examined, the propodeum is tinged with
ved at the sides, the mesosternum and the larger part of the
mesopleura are black.
Hab. North Queensland (Dodd).
ABISPA MEADE-WALDOENSIS, Sp, 0.
Almost exactly resembles in appearance the giant Odynerus,
or more probably Khynchiwm, described as Abispa par aqiuides
by Meade-Waldo, but is a true Abispa, excessively close to
A. ephippium and perhaps only a local race of this. I can see no
structural difference other than may well be individual. The
624
ON THE WASP-GENUS PARALASTOR,
whole thorax above and beneath is brownish red, the pronotum,
scutella, and propodeum laterally orange. The tomentum of the
mesonotum is pale, not black as in other species, Abdomen as
in A. ephippium. Length 24 mm.
Hab. Port Darwin (Dodd).
Fig.
COmNIOoO FP w
EXPLANATION OF THE PLATE.
. Part of Ist and 2nd abdominal segments of P. parca Sauss., in lateral
aspect. 1d. 1st dorsal segment; 2d. 2nd dorsal: 2v. 2nd ventral.
. Part of the first and the following segments of P. icarivides P. The third
and fourth segments were withdrawn within the second.
. Part of first and second abdominal segments of P. tuberculatus Sauss., in
lateral aspect.
. Median longitudinal section of the 2nd ventral segment of P. olovis in
lateral aspect (diagrammatic).
. The same of P. tricolor LP.
. The same of P. solitarius P.
The same of P. alexandrie P.
. The same of P. mutabilis P.
. The same of P. multicolor P.
. Apex of clypeus of P. imitator, very lightly emarginate. -
. The same of P. dentiger, somewhat strongly emarginate.
. The same of P. optabilis (clypeus quasi tri-dentatus).
. The.same of P. atripennis (also quasi tri-dentatus).
. The same of P. pseudochromus (clypeus fortissime emarginatus).
. The same of P. subplebeius; clypeus truncate, but slightly rounded.
. The same of P. oloris (clypeus recte truncatus).
. The five apical joints of the antenne of P. punctulatus 3.
. The small 9th, 10th and 11th antennal joints of P. hilaris, and the apical
portion of the 8th, in the terminal cavity of which the others lie.
. Three terminal joints of P. multicolor, the two latter very small and
lying in the apical cavity of the 8th.
. Tegula of P. debilis, showing the coarse puncturation of its outer portion.
. The same of P. orientalis. The coarse punctures are uot equally
numerous in all examples, but the sculpture is very different from that
of species which have the tegulz very minutely punctured.
. The same of P. alexandria. ‘The other parts of the tegul are, of course,
sculptured, but only that part (or some of that part) which gives the
best specific characters in different species is shown in this and the
preceding figures.
P. Z. S.1914, MONTAGUE, P1. I.
Bale & Danielsson, Itt
REPTILIA ETC., FROM THE MONTE BELLO ISLANDS.
‘LSAN NI SISNATIATAN SOLAYVITVH NOICGNVd DONNOA
pil ‘wossjetued y aed
Siig,
WL “Tet “SION ULINGOWNT “HALL IS) ZL
TALS vOOOMAT “Siar = tOr san Mess. Sei
oat “moOSssTeatueq W Ile
“TH el “SNOW GINOIN, “WISL -S “ZZ ‘at
GNvVaisl Hive wo NOMVLaoash ¥ NOOD vy anion
pyl ‘wossrerueqd B ee
ralicns
NEES
MU el “SOOWV INOW “HIS “SS “A “al
ON THE FAUNA OF THE MONTE BELLO ISLANDS, 625
35. A Report on the Fauna of the Monte Bello Islands.
By P. D. Monracun, B.A., Gonville and Caius College,
Cambridge *.
[Received March 18, 1914; Read June 9, 1914.]
(Plates I-IV.+)
INDEX. Pace
Moute Bello Islands: Physical; geographical .................. 625
Systematic :
WEVOTTTAITEY Sadecoconisd-cedaeos usadaanteleeeadanadaa se case uct ae ere HT3.0)
Myo RUOPSi An nOOY CES, Se Ma ase ea ea eeeeeseteesces.0ce-) O42
Risces on mdalacoonss el (Ca alate Recaniy)i esses scycee. 2. | G50
Uh OPAL OCE Ameer nessee eee eee cones cma e nc tect near eaten cet OLA!
THIGE GOCE. Gasicks Sobers codtodined doupaeatae aCe Sas E AR oe aneee =, a emetenmT of 159
Melicleptrianalovuenatcsps Ml. seen eee eee. O46
AUP OTQTE PFA OOCHHES. (0% TNS oan cceeedasbouasengucereonsesoecee ene | LOLs
WGleop erat ae tee tee esterase ce ih esate GAG
Orthop herageerantecsacecs eects eae eee ce det amends ML GAS
iv men op benam a7 praaseenes -cteme. garth teal ach same Net citer ee VGA
Wivrmayyool, — (Dir, Ireveypralbia)) » co. cocstossoeeosassobopoucerasceacan’ (eo)
Stomimamy ainGl Comets 5. s0-cceosssocccelsedososcessovensunscae, Bd)
Off the coast of Western Australia, from the North-West Cape
to Port Walcott, there stretches an archipelago of small desert
islands, the zoology of which has been but little investigated.
Lying as they do, in a shallow sandy sea, and for the most part
in sight of the low shores and mangrove-swamps of the mainland,
it is unlikely that their fauna should exhibit any marked insular
characters, but they offer great opportunities of considering the
typical forms which inhabit them in relation to their somewhat
peculiar environmental conditions. Of all these islands the
Monte Bello Group is the most isolated, and so from its position
the most likely to repay investigation.
Mr. T. H. Haynes, a gentleman engaged in experimenting
upon the artificial cultivation of Pearl Oysters, was for some
time living upon these islands, and in his leisure collected
various zoological specimens, which he sent to the British and
West Australian Museums. His house was subsequently wrecked
by a eyclone, and he was obliged to abandon his investigations.
Since many of Mr. Haynes’ specimens presented features of
interest, Mr. Bernard Woodward, Director of the West Australian
Museum, thought it desirable that the group should be thoroughly
investigated, and a grant was offered for the purpose. As this
was insufficient a special application was made to the Royal
Society, resulting in a further grant of £50. These grants
* Communicated by Prof. J. Srantey GarpineR, M.A., F.R.S., F.Z.S.
+ For explanation of the Plates see p. 652.
626 MR. P. D. MONTAGUE ON THE
enabled the writer, accompanied by Mr. L. Burns, of Perth, W.A.,
as assistant, to spend from May 29th to August 29th, 1912,
collecting upon the islands. Mr. Burns has since been drowned
while duck-shooting at the Forrest River, near Wyndham. He
was a very able collector and a delightful companion.
I must here acknowledge the very great assistance received
from Professor Stanley Gardiner at home, from Mr. Woodward
and Mr. W. B. Alexander in Western Australia, and from
Mr. Gregory Mathews and workers in various departments of
the British Museum (Natural History), who have so kindly helped
in identifying the specimens.
A detailed description of a local fauna is incomplete without
some account of its environment; it is therefore necessary to
consider the prevailing physical conditions and geographical
features of the locality before proceeding to an account of the
various forms inhabiting it.
The group lies approximately in lat. 20° 25’ S., long. 115° 30! E.,
105 miles E.N.E. of North-West Cape, and 40 miles from the
mouth of the Fortescue River—the nearest point on the main-
land. It is near this latitude that the rainfall on the West
Australian coastal region reaches its minimum, averaging under
8 inches per annum; it is of very irregular occurrence, droughts of
two or even three consecutive years being not infrequent. The
rain falls in the summer, usually in January or February, and is
frequently attended by a particularly severe form of’ eyclone
known locally as a ‘ Willi-willi.” These storms strike the West
Australian coast between latitudes 18° and 23°, and cause much
destruction to buildings and shipping. Their wind velocity has
never been accurately measured, but lines of short iron telegraph-
poles bent in the middle through as much as 45° testify to the
force attained. Cyclones may occur between November and
March; from mid-April until October, comparatively calm and
dry weather may be depended upon. During these months there
are occasional winter showers, which become more frequent and
regulay further from the coast. Hence the islands are favoured
with’ a slightly higher rainfall, and the vegetation is in conse-
quence a little more luxuriant and less scattered than that of the
mainland in the corresponding latitude. Though these showers
are refreshing when they do occur, they are really very scanty,
and contribute but little to the average, for the bulk of the rain
falls in one or more tropical downpours during the course of a
few hours.
The temperature is not excessive; during our visit the shade
temperature at mid-day averaged 82°3°, never rising above
90-0° F.
The geological formation throughout is of sandy post-tertiary
limestone, similar to that of the mainland, which extends in an
interrupted belt down the greater length of the west coast of the
continent, and is supposed to have accumulated by wind-action.
FAUNA OF THE MONTE BELLO ISLANDS. 627
The Monte Bello rock is comparatively poor in Foraminifera,
but contains numerous fragments of broken shell. Most of the
lime is concentrated near the surface, and the bare hill-tops on
Hermite Island are covered in places with an extremely dense
and hard marble-like surface deposit, only a few inches in thick-
ness, formed by rapid surface-evaporation drawing up the water
from the lower levels and depositing its mineral contents.
The main geographical features of the group are indicated on
any map. It lies at the northern extremity of an extensive
shoal, which stretches in a southerly direction to within fifteen
miles of the mouth of the Robe River, including Barrow, a com-
paratively large island lying twelve miles S. by S.W. of the
Monte Bello Group. The southern portion of this shallow area,
known as Barrow Island Shoals, has soundings of 23 fathoms,
and dries in patches at low water. This closely approaches
another bank, which lies between the mouths of the Cane and
Robe rivers, including the Mary Anne Islands. The passage
between these shoals is only four miles wide, and nowhere more
than nine fathoms in depth. Half encircling the Monte Bello
Group on the north and west sides is Breakin’s Reef, which is the
actual limit of the bank. It is nota coral formation, although
there are plenty of corals upon it, and it dries in patches at low
water with the spring tides. Outside the ‘reef? soundings of
40-50 fathoms are struck almost immediately. It seems likely
that this shallow area at one time formed an extensive triangular
cape, of which the Monte Bello Islands formed the northern
extremity, having become separated at a comparatively recent
period. Barrow Island was probably connected with the Monte
Bello Islands long after its separation from the mainland, a
supposition supported by zoological evidence hereafter to be
dealt with.
It will be weil now to describe the respective characters of the
two main islands, Hermite and Trimouille.
Hermite Island, the largest and most fertile, measures rather
over six miles from N. to S., and about one and three-quarters
across at its widest part. The coast-line is irregular, and ex-
tensive shallow inlets run far inland amongst low undulating
hills, nearly dividing the island longitudinally into an Eastern
anda Western portion. The coasts facing the open sea are rocky
and irregular, with low rugged cliffs and stony beaches; but
the shores of the inlets are regular and characteristic, the hills
sloping down to a flat rocky terrace, which is bare of vegetation
and washed by the sea at very high tides in windy weather.
This terrace descends nearly perpendicularly to the regular tidal
flat, and its edge is much undercut, owing to the tides which flow
swiftly up and down the long and narrow inlets, and to the
absenee of big seas, which would break up the configuration.
Since these inlets are locally called ‘lagoons,’ I shall continue to
speak of them as such, though they have nothing to do with the
true lagoons associated with coral formations. In the sheltered
628 MR. P. D. MONTAGUE ON THE
bays and inlets, there are growths of ‘mangrove’ comprising
an Avicennia, which grows to a considerable size and forms a
belt of varying width skirting the sandy flats Just below high-
water mark. Further out in the lagoon, the Avicennia gives place
suddenly to a true mangrove of the genus Bruguiera, which forms
another zone, and at the head of the large lagoon extends right
across to the Avicennia belt along the opposite shore.
The highest point on the island is 180 feet above sea-level.
Though the hills and ridges are low, their slopes are steep, with
much bare and weathered rock showing between the scanty
vegetation, The level plains are covered with a light, sandy, red
soil, very fertile when sufficiently watered. The vegetation is of
a type known collectively as ‘ Spinifea-serub,’ which covers large
tracts of north-western Australia. Spinifex is the predominant
plant over most areas; it grows in dense prickly tufts and
patches, in some places the shrubs so close together that walking
amongst them is a tortuous and somewhat painful business.
Next in abundance to the Spinifex is Myoporwm acununatum,
a shrub with bright green foliage and small white tlowers, very
attractive to Lycenid butterflies. An Olearia is also common ;
it is a tall and wiry shrub, with insignificant flowers, and
small and rather scanty linear leaves, giving little cover. The
branches generally bear the large cobwebs of a big and handsome
spider, Vephyla meridionalis. Of the less abundant shrubs must
be mentioned a Cassia, apparently near C. oligvelada, and a species
of Croton (Kuphorbiacez), a coarse, rough bush, generally affecting
the upper slopes of the ridges. Characteristic of the areas
along the shores may be mentioned a Chenopodiaceous plant,
Rhagodia billardieri R. Br., forming a bush of moderate pro-
portions with clusters of inconspicuous green flowers, attractive to
insects. The almost universally distributed Salsola hah, Linn.,
occurs abundantly, and Frankenia pauciflora grows commonly on
the limestone rocks, around the sheltered lagoons being seen in
isolated patches almost down to the water’s edge. Of the more
abundant herbaceous plants may be mentioned T'richolesma
seylanicum R. Br., which usually stands alone or in loose clusters
on bare patches of soil, its bright blue flowers very conspicuous.
A species of Senecio is also rather common, remarkable in that it
usually forces its way up through dense patches of Spinifex.
This brief account of the characteristic vegetation applies only
after a period of drought, such as there had been previous to our
visit to the islands. A good shower had fallen in April 1912,
but it was in all probability insufficient to bring up the majority
of the herbaceous annuals, which are short-lived, and flourish
only for a month or two after the heavy tropical rains. At such
times the island may present a different aspect.
There is no surface-water on Hermite—or in fact, on any of
the islands—during the dry season; our water-supply was
derived from a well, which had ‘been sunk for fifteen feet through
the porous rock, and which yielded us a permanent though scanty
FAUNA OF THE MONTE BELLO ISLANDS. 629
supply. After the rains, water may be obtained from surface-
diggings in the sand-hills situated to the west side of the island.
Trimouille in many respects differs markedly from Hermite
Island. The coast-line is far more regular, and there is but one
inlet. This is quite unlike the lagoons of Hermite, for there are
no rocky margins, and the dunes rise directly from shores of
white sand. It forms, in fact, a more or less circular bay, with
a narrow entrance. From the south side there runs off a tidal
creek, around which there is a considerable growth of mangrove,
but the Avicennia and Brugwera are intermixed, and do not
form separate zones. The vegetation here is also peculiar, con-
sisting almost entirely of Chenopodiaceous plants comprising
both shr ubs and herbs, mostly of the genus Atriplex, A. isatidea
Mog. covering large areas of the sandy foreshore.
This particular spot is the chief haunt of a small bird, Zon-
eginthus castanotis roebucki, which though widely distributed on
the mainland, is in the Monte Bello Group confined to Trimouille,
though individuals occasionally cross over to South-East Island,
which lies off its south-eastern end. (See p. 636.)
Nearly the whole of Trimouille is covered with blown sand,
which towards the north-western end has collected into dunes
of considerable size, two of which are particularly prominent,
the largest rising to a height of 120 feet. At the foot of this
hill, fresh water can be permanently obtained by shallow digging.
The sand at the south-eastern extremity of the island is very
coarse in texture, almost gravel, becoming gradually finer
towards the north-western end. This sifting effect, due to the
wind, was possibly of only a temporary nature, the result of a
recent gale.
As might be anticipated, the vegetation is far less luxuriant
and more scattered than on Hermite Island, though it is quite as
varied. Spinifex is much in evidence, and on the sand-hills
facing the sea one frequently meets with the widely-distributed
Tpomeca pes-capre, growing in a form with immensely long
trailing stems, with erect tufts of glossy green leaves and pink
flowers, separated by internodes as much as 20 feet in length.
Ls ribulus cistoides Linn., Boerhaavia diffusa Linn., and B. mesic
are all common, the latter often trailing up over the shrubs like
a regular climber, though it grows equally well alone.
Of the smaller islands little need be said, for they embody to
a greater or lesser extent characteristics already deseribed. There
is quite a varied flora on some of the smallest outlying islets, for
the soil has an extra fertility imparted to it by the *mutton-
birds,’ which breed in large numbers in burrows in the sand and
holes in the rocks.
In the following survey of the animals represented, stress has
been laid upon small differences which may indicate zoological
isolation, and trinomials have been adopted wherever it was
thought desirable, but only where the series show uniformity.
630 MR. P. D. MONTAGUE ON THE
Owing to the general scarcity of life, in consequence of the arid
conditions prevailing, it was often not possible to get a really
sufficient number of specimens. Particularly in the case of
insects, the matter is complicated by the fact that, where there
is no great abundance of individuals, any collected at the same
time and place are liable to be of the same brood. Thus all may
exhibit a slight peculiarity, and give a false impression of
uniformity in a character which may really be quite exceptional.
MAMMALIA.
There are now two indigenous animals inhabiting these islands,
a Hare Wallaby, Lagorchestes conspicillatus (Gould)*, and a Bat,
Liptesicus pumilus Gray. There was formerly a Bandicoot,
Isoodon barrowensis 'Thom., which until very recently inhabited
Hermite Island, but has now been exterminated.
Of introduced species, cats and black rats (J/, rattus rattus)
are numerous, and, as in other places, doing great damage
to the endemic fauna, The brown rat (J/. decwmanus) has already
established itself in a store-shed used by one of the pearling fleets,
but it does not appear to be thriving, for all the examples
observed were in a weak and diseased condition.
1. LAGoRcHEsTES consPicILLatus (Gould).
This species is closely allied to the more slender and agile
rufous-coloured “subspecies” ZL. c. leichardti, which is widely
distributed over tropical Australia. It is now confined to
Hermite and Barrow Islands, and Mr. Oldfield Thomas, who has
kindly identified the species, can detect no difference between
examples from the two localities. There is a specimen in the
British Museum, a co-type from the Gould collection, which is
recorded as having been obtained from ‘ Trimouille I., Dampier
Archipelago.” It is more than likely that this is a Monte Bello
example, for there is no island of that name in the Dampier
Archipelago, and on some of the older charts the Monte Bello
Group is represented by a single island named ‘Trimouille.
Whether it actually came from Trimouille, and not from Hermite
Island, it is impossible to decide, but at the present time the
species lives only on the latter, though old pearlers say that it
was formerly abundant on the former. The majority of known
specimens are from Barrow Island, where the species stil! occurs ;
I am aware of no other locality.
In habits it is nocturnal, hiding by day amongst the thick
Spinifex-tutts, and coming out just after sunset to feed upon the
bark and young shoots and foliage of various herbs and bushes.
It is unlikely that it will exist for many years longer, as
* The parentheses around the names of authors placed after scientific names im
this paper are used in accordance with Article 23 of the International Rules of
* Nomenclature (Proc. 7th Int. Cong. Boston, 1907, p. 44 (1912))—Eprtor.
FAUNA OF THE MONTE BELLO ISLANDS. 631
it is one of the most defenceless animals that can well be
imagined. It is easily dislodged from its hiding-place amongst
the Spinifex, from which it often rises in an awkward fashion,
tripping up and rolling over before getting away. Though it is
able to hop swiftly for a short distance, it rapidly becomes ex-
hausted, and is not difficult to obtain by simply running after
it and catching it by the tail.
The breeding:season appears to be during the summer and is
possibly dependent upon the rains, for we saw no half-grown
specimens, and as the rains of the previous summer had failed,
they may not have bred at all. It is possible, however, that the
eats had accounted for all the young ones. The pouches of all
the females were empty, but in the middle of August males and
females were often seen in pairs.
The species under consideration offers a very striking example
of “degeneration” resulting from isolation and consequent
absence of enemies. Owing to the lack of surface-water upon
these islands, it has never had to face the dingo and the
aborigines, who would make short work of an animal so easily
caught. It is remarkable that it has been able to brave the
climatic conditions for so long, and has not succumbed to a par-
ticularly severe drought or cyclone. ‘The identity of this species
with that of Barrow Island indicates a comparatively recent
land-connection, but from all the other smaller islands, which
presumably formed part of the same land-mass, the species has
disappeared.
9. IsooDON BARROWENSIS Thom.
Only shrivelled skins and bones of this species were discovered,
but skulls correspond in all essentials with specimens from Bar-
row Island, with which it is in all probability identical. The
cats have evidently been responsible for its extermination. Its
distribution corresponds exactly with that of the species last
described, and here again it is quite distinct from the corres-
ponding mainland form.
3. Epresicus PUMILUS Gray.
This little bat occurs on the North-West Australian mainland,
and is not uncommon over the whole Monte Bello Group,
appearing upon the wing about sunset. It was generally
observed near rocks and cliffs, where it probably spends the
day.
Introduced species.—It is curious to find Mus rattus particularly
common on the small outlying islands of the group which have
never been inhabited, frequenting the beaches and sand-hills
near the coast. It occurs on practically every island, and its
presence is attributed to a pearling-schooner which was wrecked
some twenty years ago. All the specimens we obtained were
632 MR. P. D. MONTAGUE ON THE
well-grown examples of typical J. rattus, with long black fur
and dark grey underparts—not the brown, light-bellied var.
alexandrinus which is more usually found in such situations.
They appeared to feed upon small crabs on the shores between
the tide-marks, and to derive their water-supply from gnawing
the stems and leaves of succulent plants. Their tracks were very
noticeable on the sand-hills, in some places forming well-worn
paths, which usually led to holes in the rocks, where they
breed.
The cats which have been introduced into Hermite Island
appear to be breeding rapidly ; wherever introduced they soon
become exceedingly shy and wary, and grow to a very large size.
They will, no doubt, in a few years time have accounted for the
wallabies, as they have for the bandicoots. If they cannot kill
a full-grown wallaby—though I am inclined to believe they do—
they make short work amongst the young ones.
AVES.
In the following notes, the nomenclature adopted is that of
Mr. Gregory Mathews, to whom I am greatly indebted for his
very kind assistance in identification. The numbers after the
names given in the notes on the different species refer to
Mathews’ *‘ Reference List of the Birds of Australia,” 1912.
LAND BIRDS.
1, GEOPELIA HUMERALIS HEADLANDI. 50 B. Pale Barred-
shouldered Dove.
Compared with the type, these birds agree in all necessary
detail. The species frequents the more bushy parts of the islands,
feeding upon fallen berries and seeds. They roost and nest in
the mangroves, constructing a loose platform of sticks amongst
the branches, upon which two white eggs are laid. The nesting-
season is dependent upon the rains, occurring usually in January
or February, but after a shower of rain on July 8th, 1912, the
males at once began to show signs of courtship, and by the end
of the month a small percentage of the birds were breeding.
Individuals were nearly always to be seen around the well and
an abandoned water-tank containing brackish water, into which
they used to fly to drink and bathe. The problem at once sug-
gests itself as to how they fared before these tanks were built.
The dews are usually heavy, and the smaller birds were some-
times observed drinking the dew-drops from the bushes in the
early morning, but, when the easterly winds are blowing, this
source fails entirely for days in succession. The only other
species which frequented artificial water-supplies is the little
“ finch,” Zoneginthus castanotis roebucki, which visited the
“ well” on Trimouille in flocks. Although at the time of our
arrival the well was choked, and several inches of material had
FAUNA OF THE MONTE BELLO ISLANDS. 633
to be removed before any water collected at all, yet the damp
sand was pitted and scored all over the surface by the beaks of
birds.
The distribution of this dove ranges from about Sharks Bay
to the mouth of the Shaw River.
3: wing 137 mm.; culmen 18; tarsus 21°.
fe) 4 | WEBI mms 7G LS 20.
Iris orange-red; bill dark slate-colour; feet salmon-pink,
claws black.
9. FIALCYON SANCTUS WESTRALASIANUS. 557. Western Sacred
Kingfisher.
Halcyon westralasianus Campbell, Emu, vol. i. p. 25, 1901.
One specimen, not fully adult, agrees with specimens from
S.W. Australia rather than with H. s. ramsayi Mathews, from
the N.W. of the continent. The specimen was shot in its haunts
among the mangroves, where it did not seem to beat all common.
@ (immature): wing 90 mm. ; culmen 36; tarsus 12.
Tris brown; bill slaty black, pale at base of lower mandible ;
feet brown, claws black.
3. Halcyon SORDIDUS MELVILLENSIS. 560A. Northern Man-
grove Kingfisher.
Mathews, Austral Avian Record, vol. 1. p. 38, 1912.
Two immature examples agree in size and general appearance
with birds from Melville Island. The specimens were obtained
amongst the thick mangroves of Hermite.
Q (immature): wing 97 mm.; culmen 49; tarsus 13.
Tris brown ; bill slaty black, pale at base of lower mandible ;
feet brown, claws black.
4. CHRYsococcyx BASALIS WYNDHAMI. 586. Western Narrow-
billed Bronze Cuckoo.
Five examples of this bird, from the unbarred immature form
to the fully adult, agree with the western bird described by
Mathews. ‘The species was found only upon Hermite, where it
is not uncommon, especially in the dense mangrove-forest at the
head of the Large Lagoon. It is a shy bird, but usually betrays
its presence by its shrill note—a clear whistle several times
repeated.
S (adult): wing 104 mm. ; culmen 12; tarsus 18.
Q (adult): ,, O(joumy ES Pep betrays
Tris dull orange-red ; culmen dark brown ; feet brown.
Young : iris grey.
5. PACHYCEPHALA RUFIVENTRIS COLLETTI. 698A. Northern
Thickhead.
Mathews, Austral Avian Record, vol. i. p. 41, 1912; Parry’s
Creek, North-West Australia.
634 MR. P. D. MONTAGUE ON THE
An immature male resembles the north-western bird described
by Mathews as above.
The specimen was shot late in June. It was in company with
several others of the same species, but, before or after that date,
no others were seen. It is probably a casual visitor to the
islands.
S (immature): wing 91 mm.; culmen 13; tarsus 20.
Iris brown; culmen light brown, dark at tip; feet dark
brown.
6. EPTHIANURA- TRICOLOR pDisTINcTA. 845. Northern Tri-
coloured Chat.
One male example has a much darker back than the type of
the above bird; darker, indeed, than any specimens examined,
but the species is variable in this respect.
The species occurs very sparingly upon Hermite; only two
examples were observed, frequenting the thick bushy localities
on the east of the island.
3: wing 69 mm.; culmen 13°5; tarsus 19°5.
Iris brown; culmen dark brown ; feet dark brown.
7. EREMIORNIS CARTERI ASSIMILIS. Island Desert Bird.
Montague, Austral Avian Record, vol. i. p. 181, 1913.
Differs from JZ. c. carteri in its smaller size, proportionately
larger bill, and in the colour of the head, which is of a deeper
and richer chestnut-brown.
The mainland representative (2. c. carteri) is a somewhat scarce
bird inhabiting the Spinzfex-country in the region of North-West
Cape. The Monte Bello form is found principally upon the
Spinifes-plains of Hermite, where it is rather common, It fre-
quents the low scrub, slipping with ease and rapidity amongst the
very thickest and most prickly bush, making goo use of its long
tail to guide and balance itself in so doing. Occasionally it will
appear for an instant at the summit of a bush or tuft, utter a
harsh clucking note, and disappear almost immediately, or make
a short and hurried flight to the next thicket.
3: wing 60 mm.; culmen 12; tarsus 19-5.
OR era comin, ae, | sob Miers
Iris brown ; culmen dark brown ; tarsus brown.
8. ARTAMUS LEUCORHYNCHUS HARTERTI. 992. Western White-
rumped Wood-Swallow.
This bird belongs to the West Australian form described by
Mathews. It was met with commonly upon all the islands,
usually to be seen hawking insects upon the wing, or, towards the
middle of the day, resting upright upon bare twigs or stumps, in
parties of three or four toa dozen. It would often accompany
us while walking over the plains, and catch insects as they were
dislodged from the Spinifex.
FAUNA OF THE MONTE BELLO ISLANDS. 635
9. ZOSTEROPS LUTEA BALSTONI, 1099. Carnarvon White-eye.
Zosterops balstoni Grant, Ibis, 1909, p. 663.
Specimens agree with the type of the above bird from Car-
narvon, North-West Australia. The species is a small, dull-
coloured form of Zosterops lutea Gould, though it would be more
correct to look upon the type lutea as an island form of the far
more widely distributed balstoni. It is much the most numerous
bird inhabiting the Monte Bello Group, living upon all the
islands, however small, where there is sufficient scrub to afford
food and protection. It appears to be omnivorous in diet, feeding
upon berries and seeds, and searching for insects amongst the
foliage of the mangroves, in the vicinity of which it is always to
be seen.
The nesting season is probably in October. In August, the
males were in full song, and at the end of that month a half-
constructed nest was discovered, suspended amongst the foliage
of a dense Druguiera, but it was not completed when I left.
3: wing 57 mm.; culmen 9:5; tarsus 17.
Oe =, OO Taming stare OR Mk Aen cal Liiy
Iris brown ; bill ath feet dark brown.
10. STIGMATOPS INDISTINCTA PERPLEXA. 1201. Allied Least
Honey-eater. :
The specimens agree with birds from mid-Westralia. On the
Monte Bello Islands the species was met with exclusively in the
larger mangrove-forests of Hermite, frequenting the densest
parts of the Brugwiera-zone. During July and August the males
were in full song. The song is particularly pleasing, s somewhat
suggestive of that of the European Reed-Warbler, but more
musical and less harsh. When singing amongst the dense and
tangled vegetation the bird is extremely difficult to locate.
3d: wing 69 mm.; culmen 14°5; tarsus 20.
Oar Oormmn: af WA Oe eee OHSe
Iris greyish brown ; bill blackish brown; feet dark brown.
11. ANTHUS AUSTRALIS MONTEBELLI, Montebello Pipit.
Montague, Austral Avian Record, vol. i. p. 181, 1913.
This is a pale subspecies, in which the dark centres of all the
feathers are much reduced, and the spotting on the breast is
comparatively sparse. It lacks any rufous tinge.
This is a common bird over the whole group, being perhaps
especially abundant upon’ Trimouille. It is the only species of
land-birds which was observed to fly from island to island over
passages more than two hundred yards across. It was met
with everywhere on the open country, both on the rocky hill-
slopes and on the sandy plains.
3: wing 89 mm.; culmen 12; tarsus 24.
Oe CONNIE TAS Sa eis aller 18
Iris brown ; bill black ; feet light brown.
Proc. Zoou. Soc.—1914, No. XLIY. 44
636 MR. P. D. MONTAGUE ON THE
12. ZoNEGINTHUS CASTANOTIS ROEBUCKI. 1347. Dark Chest-
nut-eared Finch.
This bird resembles the form from Roebuck Bay in being dark
above, and in having the dark ear-patch. The species in its
various forms is widely distributed over the Australian continent.
On the Monte Bello Group it is confined to Trimouille and South-
East Island. Its head-quarters is the lagoon towards the north-.
west end of the island, which has already been described. Large,
loose spherical nests, composed of dry herbaceous stems and lined
with feathers and soft grass, were often to be met with amongst
the branches of Avicennia, and one was found amongst the rocks
under an overhanging cliff-edge. They contained no eggs; the
breeding- season probably commences in October.
dé: wing 99 mm.; culmen 10; tarsus 14.
QS 48 Bane oe LA ie eae onae Mel bs 155
Iris orange-red ; bill orange; feet flesh-colour.
Land-birds alone are likely to furnish indications of zoological
isolation and pre-existing land-connections. It will be seen that,
with one exception, the above list only comprises birds which
inhabit the North-West of Western Australia. ‘The exception
is Halcyon sanctus westralasianus, the subspecies inhabiting
the South-Western region. The fact is of interest, as one would
naturally have expected to find H. s. ramsayi, the North-
Western bird, and the occurrence of South-Western forms on
the Monte Bello Group is found also in the invertebrate fauna.
Of the peculiar forms, Hremiornis carteri assimilis might have
been anticipated, as it is a bird addicted to one locality and
apparently incapable of prolonged flight. It has not been re-
ported from Barrow Island, however, where there occurs in its
stead a blue JdZalwrus, which is absent from the Monte Bello
Group. It is curious, on the other hand, that a bird so
widely distributed and strong on the wing as Anthus australis
should be also represented by a readily distinguished subspecies,
whereas the Zosterops, a no less characteristic and abundant
bird, of a genus quite remarkable for its insular forms, should be
identical in every respect with the type from Carnarvon.
SEA-BIRDS, WADERS, AND BIRDS OF PREY.
13. HYDROPROGNE TSCHEGRAVA STRENUA. 128. Caspian Tern.
Sylochelidon strenwus Gould, Proc. Zool. Soc. Lond., 1846.
This bird is distributed around the coasts of South and
Western Australia, and was found in scattered pairs frequenting
the low flat islets at the north of the group. A single half-
erowh young one was discovered among the rocks on a beach of
Trimouille, at the end of July.
FAUNA OF THE MONTE BELLO ISLANDS. 637
14. BRUCHIGAVIA NOVE-HOLLANDLE LONGIROsSTRIs. Silver Gull.
B. longirostris Masters, Proc. Linn. Soc. N.S. W., vol. ii.
pe flsy lsat
The range of the form extends from 8.W. to N.W. Australia.
Around the Monte Bello Islands it is not abundant ; the whole of
the north-western region is, in fact, remarkable for the comparative
scarcity of sea-birds, in spite of the existence of extensive flats and
shallows which ought to furnish an abundant food-supply.
A few old nests, apparently of this species, were found upon
Long Island and several of the outlying rocks, and the
breeding-season is said by the pearlers to be in December and
January. The nesting-time of the gulls varies enormously and
individually upon the various island groups around the West
Australian coast, occurring, it would appear, any time between
October and April.
15. H#®MATOPUS LONGIROSTRIS.
Fairly numerous around the sandy shores and mud-flats,
feeding in small parties, often in company with the next species.
16. H#MATOPUS UNICOLOR OPHTHALMICUS.
The Montebello Sooty Oystercatcher is the bird described as
above, distinguished from H. w. berniert by the bare orange-red
space around the eyes. It is really a North Australian bird,
and the Monte Bello Islands must be near the southern limit of
its range, H. w. berniert being the typical West Australian form.
17. EKupopa Georrroyi. 201. Large Sand Dottrel.
Charadrius geoffroyi Wagler, Syst. Av., Charadr. sp. 19, 1827 ;
Java.
The distribution is from Southern Siberia southwards to
Australia, where it occurs during summer, and in the northern
part of which it has often been taken in full breeding plumage.
Observed in small numbers, usually feeding in company ‘with
EH. mongolus and C. ruficupillus on the tidal flats of Hermite,
throughout June, July, and August. The specimen obtained
was shot on July 9th.
3d: wing 133mm.; culmen 23; tarsus 33.
Bill dark brown; iris brown; feet dark brown, claws black.
18. EKupropa Moncouus. 202. Mongolian Sand Dottrel.
Charadrius mongolus Pallas, Reise Russ. Reichs, vol. ii.
p. 700, 1776 ; Mongolia.
The same remarks apply as to the former species.
@: wing 129 mm.; culmen 17:5; tarsus 30.
19. CHARADRIUS RUFICAPILLUS TORMENTI Mathews. 205. Pale
Red-capped Dottrel.
Specimens agree with the above pale subspecies described by
44*
63 MR. P. D. MONTAGUE ON THE
Mathews ; it is distinct from the eastern bird. It is distributed all
along the coasts of Mid- and Western Australia and the Northern
Territory.
Observed throughout June, July, and August; being seen
generally on the tidal flats at low water, but flying about in
flocks when the tide is high along the outer shores. Specimens
were shot on June 10th and July 4th.
3: wing 102-105 mm.; culmen 13:5; tarsus 25.
20. PISOBIA MINUTA RUFICOLLIs. 230. Little Stint.
Trynga ruficollis, Pallas, Reise Russ. Reichs, vol. in. p. 700,
1776; Siberia.
The same remarks apply as to Hupoda geoffroyi and mongolus.
Specimens obtained on July 10th.
3: wing 109 mm.; culmen 17°5; tarsus 17°5.
OF VON mame oes 16; iy ee LE
91. DemreGRerrA sAcRA. Reef Heron.
This species was observed commonly, and both white and grey
forms were seen, though only the latter , which is by far the most
plentiful, was obeuned: They were generally to be found about
the rocks and cliffs of the outer aioe seldom frequenting the
more sheltered inlets. A nest, containing three eggs, was dis-
covered on a small flat islet to the north of Hermite. It formed a
lining of loose sticks to a depression in the rock, a few feet above
high-tide level. Only a yard or two away there was an Osprey’s
nest with newly-hatched chicks; although this bird will not
tolerate another nest of its own species upon the same island, it
does not in the least resent the presence of that of a different
bird.
92. PELECANUS CONSPICILLATUS. New Holland Pelican.
Observed in small parties, usually of a dozen or so, around the
islands to the north of the group. ‘They do not breed in the
vicinity.
23. HaAuiairus LeucoGAsTER Cuv. White-bellied Sea-Hagle.
G2, WE)
This species, which enjoys a wide distribution, from the coasts
of India, Ceylon, Malay Archipelago, and Australia to Western
Polynesia and Tonga, occurs abundantly on the islands off North-
Western Australia. Two pairs were nesting upon the Monte Bello
Group, one at the extreme south-eastern end of Trimouille, and
another on a small flat islet tothe north of Hermite. The former
nest was placed upon a sloping ledge of a cliff, and a great
quantity of material was employed to bring the outer edge to the
level of the inner portion ; it was consequently a bulky structure,
5 ft. 6 in. in diameter—composed of sticks and seaweed, with no
obvious cavity or lining—forming a safe and level platform for
the single chick. In the second instance, there was practically
FAUNA OF THE MONTE BELLO ISLANDS. 639
no nest, the single newly-hatched chick lying in a depression in
the flat rock, into which had been placed a scanty collection of
grass and sticks as a lining; here there was no danger of the egg
or young bird falling out and being destroyed.
The species seems to subsist entirely upon fish and sea-snakes,
never molesting the sea-birds (at least the adults), though the pair
upon Trimouille were subject to violent attacks from a pair of
Ospreys which were nesting upon a neighbouring rock. Whenever
they were in the air together, the Ospreys would circle above the
Eagle, screaming loudly, and then suddenly swoop down upon it.
The latter bird would usually turn sharply sideways or almost
upon its back, holding out its formidable talons and thus warning
off its assailants, a feat of balance and flight fascinating to watch,
24. HALIASTUR GIRRENERA Vieill.
This species occurs around the coasts of Northern and Eastern
Australia, and New Guinea. The N.W. Cape and Monte Bello
Islands are near the southern limit of its distribution on the
west side of the continent. In the region of the Ashburton
River it is not uncommon, and two pairs were observed upon the
Monte Bello Islands, where they seem to subsist almost entirely
upon small rock-crabs. On Trimouille one of these birds was
observed to leave a heap of broken limbs and empty carapaces,
which formed a pile more than a yard across upon a slab of rock in
a large shallow cave to the east of the island. A nest contain-
ing similar remains was discovered amongst the mangroves on
Hermite. It was placed in the fork of a Bruguiera, and was
deep in form, about equal in size to that of a crow and composed
of sticks, seaweed and Spinifex, with no distinct lining. It con-
tained a single egg, dirty white in ground-colour, with scanty and
minute streaks and sparks of rusty brown.
25. PANDION HALIAHTUS MELVILLENSIS. 373. Northern White-
headed Osprey. (PI. IT.)
Mathews, Austral Avian Record, vol.1. p. 34,1912; Melville I.
This Western Australian form of the almost cosmopolitan species
occurs very abundantly, particularly frequenting the archipelago of
flat rocks and islets lying to the north of the group, upon neariy
every one of which there was a nest. The majority had laid early in
June, but fresh eggs were obtained late in July and early in August.
The nest isa conspicuous and bulky structure, ranging from about
11 inches to 5 feet in height, but always measuring about 3 ft. 6 in.
in internal and about 5 feet in external diameter. It is composed
of sticks, drift-wood, seaweed and bits of coral, etc., the shallow
cavity being lined with finer seaweeds and any other soft
material. Only one nest is ever found upon an island, though
nests may be in close proximity if separated by water. When
situated upon a low rock, the nest is usually four to five feet high
and forms a conspicuous object. On the larger islands it is
640 MR. P. D. MONTAGUE ON THE
generally a smaller structure, placed in some elevated position
such as the top of a steep hill or a cliff overlooking the sea.
The eggs in nearly every case were three in number, one of
which was often addled, creamy white in ground-colour, boldly
blotched with deep reddish brown and underlying markings of
purple-grey. Average measurements, 60x47 mm. The young
when hatched are covered with grey down, and in their first
plumage the head and breast are much streaked with brown.
The food, judging from remains found in the nest, consists
mostly of sea-snakes aud a sphyreenid fish, known locally as
‘ Pike,’ which swims near the surface over the sandy shallows. In
catching prey so slippery and narrow in girth, the conical and
sharply pointed enlarged scales under the feet must be of especial
service. As has often been observed in other localities, the prey
is always carried lengthwise.
REPTILIA,
The following is a list of the Reptiles inhabiting the group.
The nomenclature adopted is that of Boulenger, Brit. Mus. Cat.
of Lizards, 1885, and Cat. of Snakes, 1893. I have to thank
Mr. Boulenger for his kind assistance in identification.
1, HEeTERoNOTA BINOEI Gray. (PI. I. figs. 1-3.)
Gray, Cat. p. 159; Boulenger, B.M. Cat. vol. i. p. 151.
This species is distributed over Western Australia and the
islands off the coast. On the Monte Bello Group it is very
abundant, and the individuals are of large size and extremely
variable in colour. It is nocturnal in habit, hiding by day under
stones or pieces of wood, but at night it may be seen in numbers
with the aid of a lantern, moving about rather sluggishly over
the sand in search of the small beetles and spiders upon which it
feeds.
2, GuHYRA vaRrEGATA, B.M. Cat. vol. i, p. 151.
Perwpia variegata Gray, Cat. p. 159.
This little gecko is widely distributed over Australia and
Polynesia, though most of the records are from the north and
west of the continent. On the Monte Bello Islands, it occurs
quite commonly on the sandy plains which are well clothed with
scrub, hiding by day in the sand, and ascending the bushes at
night in search of insects. Nearly all the examples were obtained
at night upon stumps and posts which had been ‘ sugared’ for
moths, preying upon the insects as they alighted.
The specimens obtained, which are all from Hermite, are of
small size, and show in every case four longitudinal rows of light
spots down the dorsal surface, which are absent from the majority
of examples examined from other localities. It is remarkable
FAUNA OF THE MONTE BELLO ISLANDS. 641
that on Hermite this species and //. binoei bear a strong super-
ficial resemblance to one another, both in size and coloration.
The resemblance in size is perhaps the most remarkable, as the
following figures will show :—
H, binoet: length, B.M. Catalogue, 80 mm.; Hermite speci-
mens, 103 mm.
G. variegata: length, B.M. Catalogue, 147 mm.; Hermite
specimens, 87 min.
The discrepancy between the Monte Bello forms is largely
accounted for by the long slender tail of H. binoei, the average
measurements of the largest specimens from snout to base of
tail being in the two cases, 7. binoei 58°4 mm. and G@. varregata
54mm. Furthermore, it may be observed that the white spots on
G. variegata correspond to some extent In position and appearance
to the white tetrahedral scales on the back of H/. binoei. The
fact is interesting, as the two species are found together on the
same ground, though the one obtains its food on the sand and
the other on the bushes above.
3. PHYSIGNATHUS GILBERTI. (Pl. I. figs. 4-7.)
B. M. Cat. vol. i. p. 396.
Lophognathus gilberti Gray, Cat. p. 12.
This large and handsome lizard was seen very abundantly on
Hermite Island, and occurs on nearly all the other islands. On
warm and sunny days it could be seen almost anywhere, running
and jumping with great agility over the rocks and trees, feeding
mostly upon a large and abundant grasshopper, Cyrtacanthacris
guttulosa.
The species is distributed over Northern and Western Aus-
tralia. Montebello examples are greatly below the average size.
4, ABLEPHARUS MUELLERI, B. M. Cat. vol. iii. p. 356.
Phaneropis muellert Fisch., Arch. f. Nat. 1881, p. 236, pl. xii.
This lizard is probably common, for the regular waved track,
formed by its burrowing through the surface sand, was
frequently seen in the early morning, an indication that the
species 1s nocturnal in its movements. Owing to the great
rapidity with which it burrows, it is hard to capture, and our
examples were obtained by turning over loose rocks in large
numbers. It had previously been recorded only from Western
Australia.
5. Lycosoma srpEs, B. M. Cat. vol. ii. p. 337.
Rhodona bipes Fisch., Arch. f. Nat. 1882, p. 292, pl. xvii.
‘figs. 10-15.
Apparently resembles in habit the preceding species, and
obtained from the same locality in the same manner. It is
apparently confined to N.W. Australia.
642 MR. P. D. MONTAGUE ON THE
6. Lygosoma LEsuEURII, B. M. Cat. vol. iil. p. 225.
Hinula australis Gray, Cat. p. 77.
Tiliqua australis Gray, Ann. Nat. Hist. ii. 1838, p. 291.
This species is fairly abundant on all the larger islands,
frequenting generally the limestone slopes of the hills, and during
the heat of the day running about with great activity. Distributed
over Western Australia.
7. Lygosoma 1soLepts, B. M. Cat. vol. iii. p. 234, pl. xv. fig. 1.
Hinula tenuis Gray, Cat. p. 76.
Rather common on the Spinifex-plains of Hermite. Diurnal,
feeding upon flies and the smaller Orthoptera. Distributed over
Western Australia.
8. VaRANus GouLpit, B. M. Cat. vol. 1. p. 320.
Monitor gouldi Gray, Cat. p. 12.
This large species, which occurs over Northern and Western
Australia and New Guinea, is abundant over the whole Monte Bello
Group, wherever there are flat sandy plains of sufficient extent.
In places the ground is riddled with its fattened burrows, which
usually descend to a depth of two feet or so, and ascend again to
a second entrance, though they often branch and intersect. The
reptile is only in evidence on very hot days, and is generally
shy and wary, running with great rapidity. It feeds upon the
larger Orthoptera, possibly also upon small birds. On _ the
mainland in this vicinity it is known by the name of ‘ Bung-
arra,’ derived from the aborigines, amongst whom it forms a
source of food. It is supposed to do considerable damage to
young chickens and eggs in settled localities.
9. VARANUS ACANTHURUS, B. M. Cat. vol. ii. p. 324.
Odatria ocellata Gray, Cat. p. 8.
This reptile is apparently scarce, for only one specimen was
seen, being obtained on a Spinifex-plain of Hermite. On the
mainland it is found over the north and west of the continent.
10. TyPHLops AMMoDyTES, sp. n. (PI. I. figs. 8-10.)
Snout rounded and strongly projecting; nostril lateral. Rostral
narrow, the upper portion nearly one-third the width of the head,
not extending quite to the level of the eyes. Nostril between two
nasals, the inferior of which comes just in contact with the lower
portion of the preocular. Upper nasal nearly as broad as the
rostral. Frontal small, prefrontal and parietals larger than
the scales of the body. Eyes distinct. Four upper labials.
20 scales around the middle of the body. Tail slightly longer
than broad, conical, ending in a short spine. Colour a uniform
pale greyish brown, somewhat lighter on the under surface.
Length 230mm. Diameter of body, 4mm.
Found in the sand, under a loose rock, Hermite Island.
FAUNA OF THE MONTE BELLO ISLANDS. 643
This species holds quite an isolated position. In most respects
it approaches 7’. braminis, from which it is readily distinguished
by the inferior nasal not extending to the upper surface of the
head.
11. Lrasis CHILDRENI Gray.
Gray, Zool. Miscell. 1842, p. 44; Cat. p. 93; Dum. &
Bibr. Erp. Gén. vi. p. 439, 1844.
Only one specimen of this snake was seen and obtained.
Apparently it is not common, though probably more in evidence
in the very hot weather. It is distributed over Northern Australia
to the islands in Torres Straits. J am aware of no previous
record so far south as the Monte Bello Group.
From the above list, it will be seen that the reptilian fauna is
typically North-West Australian, and it is quite possible that
the new Z'yphlops will be found also to occur on Barrow Island
and the mainland. With one or two exceptions, however, the
island forms have undergone a marked reduction in size.
The following measurements of total lengths illustrate to what
extent this reduction has taken place :—
Brit. Mus. Cat. Largest specimen obtained.
LENO UN OCU ence anes, 80 mm. 103 mm.
G. variegata ...... 147 ,, Sime
Jee OIWERGS. doenbande ANGIS) |p BOO)» oe
A. muelleri......... She AEs
Ib, WYVES hoonoocoede Bi. op 82h,
Ibe OSHA DIS saoone oon OSS WAY
Vis gould ttre 3. LOO LOMO
V. acanthurus .. 665 ,, OO
* INSECTA.
The collections of insects from the Monte Bello Islands are
small. The islands were worked only during the dry months,
when probably not one-third of the species were in the imago
state, and it is probable that after the tropical rains very
different results might have been obtained. On the other hand,
the rainfall is so erratic that there are really no fixed seasons for
emergence, and, the average temperature being high, a shower of
rain in July may cause the emergence in small numbers of a
species occurring abundantly after a heavy storm in October.
To this cause I attribute the large percentage of odd examples,
for showers occurred in April and in July, after a prolonged
period of drought—the tropical rains of the previous season having
failed. It is interesting to note, that with the common Pierid
butterfly, Belenois teutonia, though the majority were of the
dry-season form with the black on the margins of the wings
reduced, about 20 per cent. were intermediate, and a few examples
even approached the wet-season form. The collection, therefore,
644 MR. P. D. MONTAGUE ON THE
may be more representative than might at first be supposed. In
any case the number of species inhabiting the islands is likely to
be small.
LEPIDOPTERA.
Rhopalocera.
Of butterflies frequenting the Monte Bello Islands there are
eight species, the majority of which are common and _ widely
distributed forms, though one, possibly two, of the Lycnids are
distinct island forms. ‘lhe specimens have been compared with
forms in the British Museum.
1. BeLenors TevronrA Fabr.
Belenois java Waterhouse, Cat. Rhop. of Australia, no, 1
memoir, N.S.W. Naturalists’ Club, 1903.
A well-known Austro- Malayan species, very abundant.
2. PRECIS VELLIDA (Fabr.).
One example of this widely-distributed and rather variable
butterfly.
3. VANESSA KERSHAWI McCoy.
Another wideily-distributed form, the Australian representative
of the familiar V. cardui. In the Monte Bello examples, the
fifth (anterior) spot on the under surface of the hind wing is much
reduced or absent. Though in this respect the Monte Bello series
is uniform, yet these spots are generally somewhat variable, and
no systematic distinction can be made.
4. Danats curysippus Linn., form petilia (Stoll).
An Austro-Malayan form, not uncommon on Hermite.
5. NEoLuciA sERPENTATA H.-Sch.
The Monte Bello series is again very uniform, and differ from
most examples in having only one ocellus at the hind margin of
the lower surface of the hind wing. The general coloration
of the lower surface is brighter than in specimens from East and
South Australia, and less bright and distinct than in specimens
from Port Darwin and the tropical North. It is identical with
specimens from Wallaby Island in the British Museum collection,
and is possibly an island form. Very common over the whole
group.
6. NAcADUBA BIOCELLATA Felder.
This butterfly extends over Western Australia, being parti-
cularly abundant on the islands off the north-west coast. ‘The sets
obtained on Hermite, Long and Trimouille Islands are identical
with examples from Queen’s Islet, N.W.A. The species was very
abundant from June to the end of July, frequenting the flowers
of Myoporum.
FAUNA OF THE MONTE BELLO ISLANDS. 645
7. HoLocHILA HEATHII AERATA, subsp. n.
This is a common Monte Bello form of H. heathii Cox. It is
somewhat smaller, and the male shows a bronze rather than
a purple sheen upon the upper surface of the wings, and is
of generally darker coloration. In the female, the blue on the
upper surface of the fore wings is much reduced, in some examples
being confined to the base of “the interspace between the first and
second veins. Both sexes have six very distinct marginal spots
upon the lower surface of the fore and hind wings, in this
respect resembling the type and differing from examples from
the south-west and central parts of Western Australia, in which
these spots are generally indistinct and sometimes absent.
8. OXYTOXIA ARGENTEO-ORNATUS Hew.
A large series, mostly from Hermite.
Heterocera.
It is by no means sure that the majority of species in the
following list are not accidental visitors to the islands. By far
the greater number of the Noctuze were taken upon sugar on
one or two evenings, notably June 20th and from June the 29th
to the first few days of July, 1912. Though sugar was applied
almost every evening throughout our three months’ visit, yet it
was only upon these few occasions that moths were taken. The
nights in question were warm and still, following upon periods of
strong easterly and south-easterly winds, which had blown for
several daysin succession. It would not be surprising that winds
of this nature, which several times enveloped the islandsin clouds
of dust from the deserts of the mainland, should bring with them
winged insects capable of fairly prolonged flight. There are,
however, several forms which are apparently new, two of which
are described here. Other apparently new species require
comparison with type-specimens in Australia. As the N.W.
Australian fauna is at present but little known, it would be
unsafe to base any zoo-geographical conclusions upon insects so
easily transported.
1. AMSACTA MARGINATA Donov.
Three examples, similar to specimens in the British Museum
from the Sherlock River district, including a yellow-bodied
aberration, which is also identical in every respect. In the red-
bodied specimens, the black spots on the margins of the hind
wings are much reduced, whereas in the yellow-bodied specimen
they are very prominent.
2. UYETHEISA PULCHELLOIDES Hainps.
Identical with specimens from Port Darwin and Baudin
Island.
646 MR. P. D. MONTAGUE ON THE
3. Evproctris curoniris Turner.
A very abundant species distributed over most of Australia
4, AnTHELA Walk. (Darala W alk.) PUDICA Swinh.
A somewhat dark form of this West Australian species.
5. CoLtusa sp. ? near C. flavala.
6. EUxoA RADIANS.
Distributed all over Australia.
7. CHLORIDEA ARMIGERA Hiibn.
Agrees with specimens from North Australia, in which the
orbital stigma is nearly obsolete.
8. NEOCLEPTRIA PUNCTIFERA Walk.
Distributed over Northern and Western Australia.
9. MELICLEPTRIA NEURIAS Meyr.
Agrees with the light West Australian form from the Sherlock
River.
10. MELICLEPTRIA ALBIVENATA, sp. n. (PI. I. fig. 11.)
Head and thorax light brown ; abdomen white below, yellowish
above. Fore wings yellowish brown, the veins white, narrowly
bordered on each side with dark brown. In some specimens, the
wing more or less diffused with scattered white scales, with the
exception of the diseoidal cell, which stands out rather clearly,
and the region near the outer margin. A terminal series of dark
points, but often indistinct. Hind wings white, suffused with
brown except upon the terminal and costal areas. Expanse,
27 mm.
In general appearance resembles J/. canusina Swinh. (Hamps.,
B.M, Cat, vol, iv. p. 99) from the Sherlock River district, but
distinguished by the white nervures and suffusion.
Taken on sugar, Hermite, July 1912.
11. Proparria MUNDOoIDES Lower.
Widely distributed upon the mainland of Australia.
12. EcropaTRIA ASPERA Walk.
Distributed over Australia and New Zealand.
13. PANDESMA SUBMURINA Walk.
Australia and New Guinea.
14. Grammopus ocELLATA Tepper.
Two typical examples. Distributed over Australia.
15. GoNITIS SUBULIFERA Guen.
A few examples of this almost cosmopolitan species.
FAUNA OF THE MONTE BELLO ISLANDS, 647
16. ANUMETA ZUBOIDES, sp. n. (PI. I. fig. 12.)
This may prove to be merely a form of Anwmeta zuba Swinh.,
to which in any case it is very closely allied. Compared with
specimens in the British Museum, the following distinctions are
readily observed : somewhat larger in size; the white line across
the fore wings in the majority of specimens much more distinct,
waved instead of straight ; hind wings darker in colour. Expanse,
50 mm.
The specimens were taken on sugar, Hermite, July 1912.
17. CIRPHIS ABDOMINALIS Walk.
Recorded from North Australia. Hermite, scarce.
18. Amywna ocro Guen.
Widely distributed, occurring from India to the New Hebrides.
19. Amyna sprtonota Lower.
A North Australian species, recorded from Port Darwin.
20. EUBLEMMA DUBIA Butler.
Distributed over the greater part of the Australian continent.
21. PotypEsMA LAwsont Feld.
Western Australia. Specimens identical with examples from
the Sherlock River.
PoOLYDESMA MARMARINOPA.
Also identical with specimens from the Sherlock River.
COLEOPTERA.
The following very meagre provisional list comprises all the
beetles found upon the islands, although considerable pains were
taken in their collection. Searching the sand and vegetation at
night by the aid of a lantern yielded the best results, but there
ean be few species and those by no means abundant. Of these
three appear to be new, but they are not now described, as many
type-specimens of West Australian species have not been examined.
The list will serve at least to show the type of the fauna.
Trox crotchi Har.. Hermite.
. Isodon novitius Blackb. Hermite.
. Coccinella transversalis Fabry. Hermite.
Dermestes cadavorinus Faby. From a dead Cat.
. Gonocephalum meyricki Blackb. Hermite.
. Sympetes, (2?) sp. n. Hermite.
. Saragus, (?) sp.n. Hermite.
. Mictotragus arachne Pascoe. All islands.
. Phorocantha, (2) senio Newm. Hermite.
Symphyletes, (2) sp.n. Hermite.
Bostrychopsis jesuitus Fabr. All islands.
ow oo bo Ee
is
HOw naASD
648 MR. P. D. MONTAGUE ON THE
ORTHOPTERA.
The Orthoptera, as is usually the case in warm and arid
localities, comprise a large and characteristic section of the insect
fauna of the Monte Bello Islands. There are few species, but of
those that do occur, some are extremely abundant. The larger
winged species are mostly of wide distribution, but the smaller
wingless forms comprise several undescribed (¢) species.
The aeridiids include such forms as the large Cyrtacanthus
guttulosa, which occurs in great numbers upon all the islands,
forming the main food-supply of the larger Reptiles. This insect
is capable of prolonged flight, though it is only upon rare occasious
that it exerts this faculty, and then can be observed flying in
swarms through the Spinifex, usually about sunset. The ‘ Blue-
winged Locust,’ Coryphistes cyanopterus Charpentier, also occurred
but the examples were mostly of small size. The species is of
wide distribution. Another large form, Acridiwm maculicollis,
was also recorded, though it was not common.
Of the mantids, only two were met with in numbers,
Archimantis brunneriana Sauss., a North Australian species, and
a smaller species of the genus Orthoderides, near O. ministralis
Fabr., but as yet undetermined. This insect was always found
upon the foliage of Wyoporum acuminatum, the leaves of which it
closely assimilates both in form and colour.
Three species of cockroach were obtained, two of them identical
with, or closely allied to, Periplaneta concolor Walk. and
Ellipsidron inquinata Walk., and the third apparently a new
species of the genus Polyzosteria. One female phasmid was
also obtained, clearly a form of or closely allied to Wyrtacus
entrachelia (Westwood).
HYMENOPTERA.
The species of Hymenoptera from N.W. Australia are little
known. Consequently any geographical conclusions based on
collections from Monte Bello are likely to be erroneous. The
following is a provisional list showing the type of the fauna, but
the ants have been omitted.
1. PARACOLLETES PERFASCIATUS Cock.
This species has been identified by Mr. G. Meade-Waldo, British
Museum, who contributes the following note.
“The type of this species, described by Cockerell (Ann. Mag.
Nat. Hist. (7) xvii. p. 25, 1906) from a specimen in very pocr
condition, has ‘Western Australia’ as its only locality. It may be
useful to give a few remarks on the pubescence, which is
much spoiled by wetting in the type. For structure, Cockerell’s
description is excellent. The thorax in fine fresh specimens is
densely clothed with brownish-buff pubescence, thickly inter-
FAUNA OF THE MONTE BELLO ISLANDS. 649
mingled with black hairs; the median segment is clothed with
whitish pubescence on the surface of the truncation, and the scopa
on the posterior tibie and tarsi is composed of long and dense white
hairs. Two females collected on Hermite Island, July 1912.”
2. NoMIA FLAVIVIRIDIS, subsp. doddi, Cock.
Also identified by Mr. Meade-Waldo. Another W. Australian
species.
. Epactrothynnus productus Turner. Hermite.
. Thynnus, (?) sp. n. Hermite.
. Bembex, (2) variabilis Sm. Hermite.
. Sphex australis Sauss. Hermite.
. Trachysphex, (2) pilosulus Turner.
. Saluis tuberculatus Sm. Hermite.
. Aporus cingulatus Fabr.
. Aporus, (?) sp. n. Hermite.
11. Ephutomorpha morosa Westw. Hermite.
12. Ephutomorpha modesta Smith. Hermite, Trimouille.
13. Henicospilus sp. ?
Oo MIS Oe Co
=
Of other insects the Hemiptera were not well represented, the
majority of forms being Homoptera. Two species of water-bugs
were obtained from the Home Lagoon, Hermite, one closely
resembling Hermatobatodes marche: Coutiére et Martin (Bull,
Mus. Hist. Nat. Paris, vii. 1907, p. 214) and the other Halobates
wulterstorfii (Frauenf.)* ; both appear to have points of specific
difference.
MYRIAPODA.
Identified by Professor K. Kraepelin.
The species are all forms found upon the West Australian
mainland, though one is apparently an insular variety. The
specimens were largely collected from under stones during the
daytime, but some were found running about in the sand by the
aid of a lantern at night.
1. RHOMBOCEPHALUS L&TUS Haase.
Scolopendra leta Haase, Abh. Mus. Dresden, No. 5, p. 51,
pl. iii. fig. 51 (1887).
Recorded from Western Australia and New South Wales.
Fairly numerous on Hermite.
2. RHoMBOCEPHALUS MoRSITANS (Linn.).
(Scolopendra morsitans Linn.)
Probably imported from Asia by way of the mainland, where
it is now established. ‘The species is now almost universally
distributed in the warmer parts of the world.
* Ofr. F. B. White, ‘ Challenger’ Report, p. 40,
650 MR. P. D. MONTAGUE ON THE
3. CoRMOCEPHALUS TURNERI Pocock.
Pocock, Annals & Mag. Nat. Hist., 7th Series, vol. viii. p. 456
(1901).
Three examples, taken on Hermite, differ from the type from
Perth, Western Australia, in that the last tergum has a distinct
median groove, which is lacking in the mainland form.
4, ASANADOPSIS MJOBERGI Kraepelin.
At the time of writing, Professor Kraepelin’s description of this
species, which was sent to him by Dr. Mjéberg from North-
Western Australia, was not yet published. Professor Kraepelin
considers this second example from Hermite to be identical,
though he could not be absolutely certain, for he had sent back
the type and was consequently unable to compare the two.
One species of scorpion was obtained on Hermite. It is
identified by Dr. Kraepelin as Lychas variatus Thor.
PISCES.
The following fishes are inhabitants of the tidal lagoons. They
nave been identified by Mr. C. Tate Regan.
1. Orectolobus tentaculatus Peters.
Rhinobatus armatus Gray.
. Murena thyrsoidea Richards.
Plotosus anguillaris Bloch.
Cheroichthys valenciennei Kaup.
Pseudochromis fuscus Miller & Trosch.
Valenciennea longipinnis Benn.
. Gobius phalena Cuy. & Val.
. Opsanus diemensis Lesueur.
oe wt
Io
woe 2)
SUMMARY AND CONCLUSIONS.
In considering the relation between these islands and the main-
land, a feature of primary importance is the extreme shallowness
of the sea between them and the mainland, in all probability
indicating a separation of comparatively recent date. The
invertebrate fauna of Barrow Island is unfortunately little known.
The identity of the Wallaby, Lagorchestes conspicillatus, and the
Bandicoot, /soodon barrowensis, support the theory that Barrow
and the Monte Bello Group were continuous long after their
separation from the mainland.
The fauna, however, is not typical Vorth-West Australian, such
as is met with in the Ashburton district, but comprises many
Northern and South-Western forms. There are, furthermore, few
species, and these are of mixed distribution. Many characteristic
North-Western forms are entirely absent, though a high per-
centage of those species not. easily transported (such as the
Reptiles and Chilopoda) are identical with, or slightly modified
representatives of, the species occurring in the immediate vicinity
upon the mainland. Mr. Hogg, in dealing with the spiders, has
FAUNA OF THE MONTE BELLO ISLANDS. 651
pointed out the absence from the collections of several species so
widely spread as to be generally found in any locality in Australia.
He suggests, therefore, that the islands have once been largely
denuded of the original spider fauna, and were repopulated
subsequently with other species. These seem mostly to have come
from the south-west by the trade-wind course, for the majority
could certainly have been wind-borne. ‘These remarks in some
measure seem to apply to other groups of animals, though many
species are northern, occurring in Queensland and the Northern
Territory, probably having come down by the hurricane course.
That an almost complete depopulation of the islands could have
occurred is by no means an improbability, when their small size
and the prevailing meteorological conditions are considered. It
will be noted that the vast majority of the specimens recorded are
from Hermite. ‘This island was certainly the most thoroughly
worked, though the others were frequently visited, but its fauna
was at least three times as large as that of any of the smaller
islands. This is merely a demonstration of the obvious fact that
the smaller the island, the less will be the chance of a sufficient
number of individuals surviving to perpetuate the species after a
particularly severe and prolonged period of drought. It is indeed
remarkable that the Wallaby has been able to survive, considering
that it lives entirely in the open, and has nowhere to shelter
except amongst the Spinifex during the hurricanes, when trees
and bushes are torn up by their roots, and sand and shells blown
about with such violence as to cut deeply into wood.
In considering this point, it 1s well to remember that little work
has been done in the North-West of Australia, and the distribution
of some species may be found to be much more extended than is
at present supposed. Moreover, other forms, which might be
looked upon as island species, may really be quite common on the
mainland, though as yet unrecorded. However, the well-known
generalisations with regard to island faunas ( using the word
‘island’ in the zoological sense) hold good:—the wingless forms
not easily transported comprise most of the undescribed forms,
being In many instances clearly the modified representatives of
mainland species, whereas the easily transported winged species
are for the most part of wide distribution. There are many
anomalies, which can only be explained by considering individually
the habits of each species and the prevailing meteorological
conditions. Amongst the Longicorn beetles, for example, there
are two abundant species, of which one, Mictotragus arachne, has
the elytra completely fused, and is absolutely identical with the
form occurring abundantly upon the mainland opposite. The
other, Symphyletes, sp.n.(?), a form with well-developed wings, is
confined to Hermite. A possible factor in such local distributions
is perhaps to be looked for in the whirlwinds which area character-
istic weather-feature of the district. These disturbances can be
observed almost any day, and must certainly be regarded as an
important distributing factor. Of the two beetles in question, the
Proc, Zoou, Soc.—1914, No, XLY, 49
652 ON THE FAUNA OF THE MONTE BELLO ISLANDS.
Mictotragus is almost invariably found walking about freely upon
the surface of the sand, and has an extremely thick and hard
cuticle, being therefore well adapted for wind transportation.
The Symphyletes, on the other hand, was only found upon its
particular plant, Zricholesma zeylanicum R. Br., clinging to the
stems just below the clusters of flowers and buds, from which it
was not easily dislodged.
Of the birds, one new subspecies, Hremiornis cartert assimilis,
is a bird of very weak flight, seldom fluttering more than a few
yards at a stretch, and taking shelter in the densest thickets. It
is therefore not surprising that it should differ from its mainland
representative. In the subspecies of Anthus australis, however,
the case is different, for it is a bird of powerful flight, often
observed crossing the passages between islands, and inhabiting the
open plains. It is suprising that this bird should exhibit even
more marked differences from the mainland form than the last-
named,
EXPLANATION OF THE PLATES.
Prats I.
Heteronota hinoei.
Figs. 1,
. Showing variation in markings.
Individual with regenerating tail.
ws)
Physignathus gilberti.
. Male, nat. size.
. Under surface of head, showing general outline and gular fold,
. Female, nat. size.
. Under surface of head of female.
TAA
Typhlops ammodytes, sp. n.
8. Head, dorsal view.
9. Head, ventral view.
0. Tail, ventral view.
11. Melicleptria albivenata, sp. n., nat. size.
12. Anumeta zuboides, sp. n.
Prate II.
Young Pandion haliaétus melvillensis in nest.
Prare III.
Nest and young of Haliaétus leucogaster.
Prate LV.
Home Lagoon and Vegetation of Hermite Island.
PZ.S.1914.RATHBUN.PLI,
Huth, London.
CRUSTACEA FROM THE MONTE BELLO ISLANDS.
PZ.8.1914.RATHBUN PLU
Huth,London,
CRUSTACEA FROM THE MONTE BEL nO- tS Anieee
ON CRUSTACEA FROM THE MONTE BELLO ISLANDS. 653
36. Stalk-eyed Crustaceans collected at the Monte Bello
Islands. By Mary J. Ratrusun, United States
National Museum, Washington, D.C., U.S.A.*
[ Received March 18, 1914: Read June 9, 1914.]
(Plates I., II.+)
InpDEx.
Systematic : Page
Periclimenes hermitewsis, Sp. We... ...ceceeecsevseeenserreveee 650
fiaitknarates WES Dery INOVNS Ts“ pocenseencca casos bnonoaesaeeqesnces HSK /
AIGTED: GUS ARLy FD Ws oop coscovsvcncondeneccassceanoecseonnc oats)
Glyptowanthus cymbifer, SP. MN. ... ccc see eee seeseeseesrsensese 858
This little collection, numbering only 28 species, contains three
new species and variations of several old ones. Incidentally a
hew name is given to one of the numerous forms of Zhalamita
related to 7. admete, which in time ean perhaps be lined up as
subspecies or varieties of that species. ‘The most notable of the
new species is the Glyptoxanthus, a genus new to the Indo-Pacific
region.
A remarkable discovery is the fact that at least one of the
marine crabs, Vasxioides serpulifera, undergoes transformation to
the adult form while it is still within the breod-pouch of the
mother. Furthermore, it there passes through two adult stages,
but whether the first is hatched directly from the egg or not, it
is impossible to tell. It has long been known that the fresh-
water crabs (Potamonide) develop into the adult form before
leaving the mother, and in some cases at least pass through a
megalops stage. Careful researches into the metamorphoses of
marine crabs would doubtless disclose other cases similar to that
here recorded.
Order DECAPODA.
Family PENEID A.
METAPENEUS MoNOCEROS (Fabricius) £.
Metapeneus monoceros Alcock, Catal. Ind. Dee. Crust. Ind.
Mus., Part iii. Fasc. i. 1906, p. 18, pl. iii. figs. 7—7e.
Dredged off Hermite; 2 fathoms; July 7, 1912; No. 110:
1 S juv., about 47 mm. long.
METAPENEUS STRIDULANS Alcock 2
Metapeneus stridulans Aleock, Catal. Ind. Dec. Crust. Ind.
‘Mus., Part: iii. Fase. i. 1906, p. 27, pl. v. figs. 14, 14 a-d, and
synonymy.
* Communicated by Prof. J. Srannny Garprner, M.A., F.R.S., F.Z.S.
+ For explanation of the Plates see p. 664.
{ [The parentheses around the names of authors placed after scientific names in
this paper are used in accordance with Article 23 of the International Rules of
Nomenclature (Proc. 7th Int. Cong. Boston, 1907, p. 44 Gore) pat Dinos
45
654 MISS M. J. RATHBUN ON CRUSTACEA
Monte Bello Islands ; special locality not given: 19.
In the main, this specimen agrees with Alcock’s description
and figures. There are 17 ridges in the stridulating organ near
he posterior corners of the carapace. The telson and the
helyeum differ from typical stridulans. The telson has 5 (not 4)
pairs of marginal spines, 4 pairs of which are movable, and the
posterior pair immovable. The thelycum has the plate between
the legs of the fifth pair similar to that in fig. 14d, loc. eit.; the
plate between the legs of the fourth pair is trilobate or in the
shape of a clover-leaf; the bar between those two plates is longer
(from front to back) and narrower than the same plate in
Alcock’s figure.
On p..50 of his Catalogue (op. cit.), Dr. Alcock suggests the
identity of my akayebi with stridulans. M. akayebi, however, is
quite distinct ; the body and rostrum are more slender, the teeth
on the latter more prominent; the antennal, hepatic and branchi-
ostegal spines are larger; the second and third segments of the
antennular peduncle are longer and more slender; subterminal
prominence on carina of fifth abdominal segment obtusangular,
not dentiform; sixth segment longer and narrower than in
stridulans. The thelyecum is similar to that of the female from
the Monte Bello Islands.
Family CRANGONID& (=Alpheide),
CRANGON EDWARDsII (Audouin).
Alpheus edwardsii de Man, Jour. Linn. Soe. London, Zool.,
vol. xxii. 1888, p. 266, and synonymy.
Home Lagoon, Hermite; Aug. 12, 1912; No. 140: 1 9 ovig.,
51 mm. long.
Monte Bello Islands; special locality not given ; 2 specimens;
the larger one lacks both pairs of chelipeds, the smaller one lacks
the larger cheliped of the first pair.
The two specimens possessing chelipeds of the second pair
belong to the variety in which the first and second joints of the
carpus are of the same length. The larger of these specimens
shows a faint notch in the upper margin of the smaller manus of
the first pair.
CRANGON BUCEPHALUS (Coutiere), var.
Alpheus bucephalus Coutiere, Alpheide in Gardiner’s Mald. &
Laccad. Archip., 1905, p. 890, pl. Ixxviii. figs. 29-297; De Man,
Alpheide ‘ Siboga,’ 1911, p. 316.
Dredged off Hermite; sandy bottom; July 9-12, 1912; No.
109: 13, about 12°5 mm. long.
The differences between our specimen and the type are so few
that the former probably represents no more than a variety of
the latter. The specimen in hand is considerably larger than the
FROM THE MONTE BELLO ISLANDS. 655
type; it has not so longa rostral tooth, and consequently the
sinuses between that tooth and the orbital prominences are not
so deep; the antennal peduncle is not so long as in Coutiére’s
fig. 29, although it is a little longer than the antennular peduncle ;
the merus of the larger cheliped has a tooth at the lower distal
angle, not unlike the tooth shown in Coutiére’s fig. 29 a’, but he
says (p. 821) “le méropodite est inerme,” and in the figure, the
tooth is applied against the carpus so that it is inconspicuous.
Family PALHMONIDS®.
Subfamily Pontoniine.
PERICLIMENES HERMITENSIS, sp. n. (PI. I. figs. 1-3.)
Hermite; under rock; Aug. 17, 1912; No. 159, 1 2 ovig.
Dimensions. Type @, length from: tip of rostrum to tip of
telson, approx., 39 mm.; length of carapace, 14 mm.
A Perielimenes with thorax strongly arched from front to back
as well as from side to side. Rostrum reaching to middle of
third segment of antennular peduncle, compressed, thin; upper
margin convex, armed with six teeth, the posterior of which is
slightly behind the orbit ; tip acute ; lower limb shallower than
upper, margin slightly convex, unidentate. An antennal and an
hepatic spine present; no supraorbital spine. Branchiostegal
angle of carapace rounded. Eyes small. Seale of antennulee
narrow-ovate, reaching nearly to end of second segment, basal
spine curved, acuminate. Peduncle of antenne reaches just to
end of first segment of antennul, basal spine small; scale ovate-
oblong, most produced at inner angle, its outer spine small, scale
reaching beyond the peduncle of the antennule but not so far as
their flagella; the flagellum of the antenne when bent back
reaches half the length of the abdomen.
The added lengths of the last two articles of the third maxilliped
exceed that of the antepenultimate article. The chelipeds of the
first pair when extended reach beyond the acicle by the length of
their chelze ; merusand carpus subequal, manus 13 times as long as
fingers and 2 as long as carpus. Chelipeds of second pair sub-
equal, reaching beyond the acicle by the length of the chele ;
merus cylindrical, unarmed; carpus cup-form, a large V-shaped
notch in the lower, outer portion of the distal margin; manus
subcylindrical, gradually widening a little distally ; fingers nearly
as long as palm, prehensile edges entire and meeting or over-
lapping when closed except at the base, where there is a very low
tooth on the fixed finger between two similar ones on the dactylus ;
tips spiniform, turned sharply toward each other.
The dactyli of the third, fourth and fifth feet are short
(about 1 as long as the propodi), curved ; lower margin convex at
the base, but without spines or other protuberance.
The first three segments of the abdomen are very broad ; sixth
segment not much longer than wide and 2 as long as the telson ;
656 MISS M. J. RATHBUN ON CRUSTACEA
the latter is triangulate, with nearly straight sides, and an ob-
tuse angled tip armed with three pairs of spinules, of which the
intermediate pair is longest and the outer pair shortest. Uropods
broad-oval, longer than the telson, the outer pair broader and a
very little longer than the inner pair.
Colour. 'The specimen, which is preserved in formalin and gly-
cerine, is handsomely marked with lines of crimson and purple 3
on the pleura of the first three abdominal segments there are four
incomplete oval areas partly outlined with narrow crimson stripes ;
the same colour forms aring on the end of each uropod, the inner
ring smaller than the outer; the tip of the telson is also outlined
in the same colour; within these five areas of the tail-fan, the
colour is a lighter yellowish-red, Dorsal face of carapace a brown-
ish-orange. Chelipeds of second pair with a narrow stripe of
purple at the distal end of the merus, carpus and manus, and «
more crimson stripe on the distal half of the fingers. The third,
fourth, and fifth feet have similar purple stripes on the three
principal segments.
This species is not very nearly related to any other. It has
the same rostral formula, . as P. parvus Borradaile (see Willey,
Zool. Results, Part iv. 1900, p. 407, pl. xxxvi. figs. 3-3), but
that species is more slender, with lower rostrum and larger eyes.
ANCHISTUS INERMIS (Miers).
See Borradaile, Ann. Mag. Nat. Hist. (7) vol. ii, 1898, p. 387.
Hermite, in mantle-cavity of Pinna; July 7, 1912; No. 112:
19 ovig., about 26 mm. long.
This corresponds in the main with the description and figures
given by Miers (Crust. ‘ Alert,’ 1884, p. 291, pl. xxxii. figs. B, b,
b'). In the Hermite specimen, the telson has at the end four
sete, the outer pair stouter than the slender inner pair but nearly
as long; Miers’s fig. b shows five setze, one median and two lateral
pairs, the outer pair about half as long as the inner. The smaller
cheliped of the second pair in our specimen has the tooth on the
dactylus feebly developed, and fitting into a cavity in the fixed
finger; in Miers’s'type this tooth was not developed, but the
specimen was smaller.
Miers described the species from Port Molle, Queensland ; he
also had a specimen taken from a Pinna at Sharks Bay, West
Australia.
Family PagURID 4.
DarpANus MEGISTOS (Herbst).
Pagurus punctulatus Aleock, Catal. Ind. Dec. Crust. Ind. Mus.,
Part il. Fase. i. 1905, p. 81, pl. viii. fig. 1.
One specimen, of medium size. he species is distributed
throughout the Indo-Pacitfie region.
‘FROM THE MONTE BELLO ISLANDS, “657
Family PorRTUNIDA.
THALAMITA DISPAR, hom. n. (PI. I. fig. 4.)
Thalamita savignyt de Man (not A. Milne-Edwards), Zool.
Jahrb., Syst., vol. viii. 1895, p. 564.
Monte Bello Islands; 3¢.
Dimensions. Type and largest 3: length of carapace 15:2 mm.;
width 25°4 mm.
This is the same species as that described by de Man (oe. cit.),
for I have at hand for comparison one of his specimens from
Palos Bay, West Celebes ; but it cannot be the true savignyi of
A. Milne-Edwards (Arch. Mus. Hist. Nat. Paris, vol. x. 1861,
p- 357), because in the figure of this in Savigny’s ‘Egypte’ (Crust.
pl. iv. fig. 4) the inside of the hand shows coarse granules.
Carapace pubescent except on the elevated ridges. These are
prominent; the posterior one across the cardiac region with a
smal] piece on the branchial region, though bare and elevated,
lacks the distinct granulated edge which characterizes the other
ridges. Lobes of front separated by an open median slit. Fourth
tooth of lateral margins absent. Five crests on upper and outer
surface of palm, the two uppermost crests each with three spines,
of which the terminal one is short and subacute; third crest
tuberculate, fourth and fifth obscurely granulate, the fifth very
finely so; the first three interspaces each with some coarse
granules and also pubescent ; in the smallest of the three speci-
mens, the third interspace is smooth; there may be a line of
pubescence above the lowest crest but it is not constant.
The above characters suffice to differentiate this from other
members of the admete group.
The Celebes specimen is an adult female smaller than any of the
males. It differs from them in having a very shallow and small
median emargination in the front. There is a minute rudiment
of a fourth lateral tooth on the right side only.
Family XANTHID&.
CARPILODES RUBER A. Milne- Edwards. i
Carpilodes ruber A. Milne-Edwards, Nouv. Arch. Mus. Hist.
Nat. Paris, vol. i. 1866, p. 228, pl. xi. figs. 4-40. ;
Without special locality ; 2 9 immature, 13 mm. and 14°4 mm.
wide. They retain a deep crimson colour in alcohol.
Home Lagoon, Hermite ; under stones; July and August, 1912;
No. 108: 3 2 (1 mature, 2 young).
ATERGATIS OCYROE (Herbst).
Atergatis floridus Aleock, Jour. Asiat. Soc. Bengal, vol. xvii.
1898, p. 98.
Without special locality: 1 g,1 2 juv.
858 MISS M. J. RATHBUN ON CRUSTACEA
PLATYPODIA GRANULOSA (Riippell).
Lophactea granulosa Alcock, Jour. Asiat. Soc. Bengal, vol.
Ixvii. 1898, p. 101.
Without special locality: 3 ¢,2 9,2 juv.
ACTA AFFINIS (Dana).
Acteodes affinis Dana, Crust. U. 8. Expl. Exped., vol. i. 1852,
p- 197; Atlas, 1855, pl. xi. fig. 3.
Home Lagoon, Hermite; under stones; July 1912; No. 105:
Se:
ACT#HA GLANDIFERA, sp.n. (PI. I. fig. 5.)
Type-locality. Monte Bello Islands: special locality not given ;
26,1 2 (1 @ is type).
Additional locality. Home Lagoon, Hermite, under stones ; July
OZ aN On Oil aL Os.
Dimensions. Type 3g, length 8:4 mm., width 12 mm,
Closely related to Actea spinosissima Borradaile*, from which
it differs in the character of its tubercles and spines rather than
in their position. The tubercles of the dorsal surface are broad
and arcuate as in spinosissima, but are thicker and less flattened
and less petaloid, those toward the lateral margins being acorn-
shaped. This is also the form of the five antero-lateral spines
behind the orbit; in spinosissima they are long spines. The
projections of the front are simply crenulations, not spines nor
spiniform teeth. Spines of chelipeds short, stout, acorn-shaped,
and directed a little forward as are those on the carapace. The
spines on the legs are longer than on the chelipeds and are stout
and subacute, not slender and elongate as in spinosissima.
Although the five specimens are all larger than any specimen
of spinosissima yet recorded, there is no indication of intergrading
from one species to the other.
GLYPTOXANTHUS CYMBIFER, sp.n. (Pls. J., I. figs. 6, 7.)
Type-locality. Monte Bello Islands ; no special locality given :
2 3 (lis type), 1 9.
Dimensions. Type $, length of carapace 10°6 mm., width of
same 17mm. Paratype 9, length of carapace 12°7 mm., width
of same 20 mm.
The carapace is closely covered with small bead granules, and
is deeply areolated; the protogastric regions are divided into two
oblong lobules by a longitudinal furrow; the branchial region
bears about six lobules of irregular size, the two next the inner
angle of the region being connected by a posterior elevation.
There are four small tuberculiform teeth on the lateral margins
* In Gardiner, Fauna Mald. & Laccad. Arch. i. Part 3, 1902, p. 256, text-tig. 53.
FROM THE MONTE BELLO ISLANDS. 659
behind the angle of the orbit; they are widely separated by a
granular rim ; all these teeth except the posterior one are above
the true margin, which inclines downward toward the angle of
the buccal cavity, as in all the members of this genus. The
frontal lobes are obliquely truncate, and separated by a broad,
shallow V.
The upper-outer surface of the carpus and the upper surface
of the manus are occupied by deep, more or less rounded cavities,
with rims which are microscopically granulated but appear smooth
in relation to the general granulation of the body. The blackish-
brown colour of the fingers runs far back on the lower part of the
palm in the male, but is confined to the digits in the female;
their granules are set in a felt-like background.
The ambulatory legs are remarkable in having the upper
surface of the carpal and propodal segments each occupied by a
deep cavity bordered on the posterior margin by a thin rim,
somewhat resembling a longitudinal section of a serpulid tube,
and on the anterior margin by a thick band of pointed granules
irregularly placed.
Lower surface of crab hairy, the hairs of two kinds, one fine,
the other coarse, but both soft.
Glyptoxanthus has hitherto been known only from the coasts
of middle America, West Africa, and the Cape Verde Islands.
G. erosus (Stimpson) from the West Indian region attains a
width of about 4em. When small, about the size of these spe-
cimens of G. cymbifer, it has a granulated surface; with age
the granules wear down smooth, giving it a much more eroded
appearance.
Our species differs from all the other described species in the
curious hollows or cups on the chelipeds and legs.
XANTHIAS ATROMANUS (Haswell).
Aanthodes atromanus Haswell, Catal. Austral. Crust. 1882,
joe AS), alle ie cies, I
Home Lagoon, Hermite; June 1912; No. 106: 3 2 ovig.,
varying from 8°7 mm. to 13°6 mm. in width.
PHYMODIUS UNGULATUS (Milne-Edwards).
For variations, see Phymodius wngulatus Rathbun, Mem. Mus. Comp. Zool.
vol. xxxv. 1907, p. 46, pls. iti. & iv.
Monte Bello Islands: 1 9 immature.
Length of carapace 8 mm., width 11 mm.; width of front, orbital
angle excluded, 4 mm.
In this specimen the areoles of the carapace are very much
subdivided, the lateral marginal lobules obtuse except the last
which has a very short spinule at the tip, the chelipeds are sub-
equal, the palms have subparallel margins and acute tubercles,
the fingers are slightly curved and have a narrow gape.
660 MISS M. J. RATHBUN ON CRUSTACEA
PILUMNUS VESPERTILIO (Fabricius).
Pilumnus vespertilio Alcock, Jour. Asiat. Soc. Bengal, vol. Ixvii.
1898, p. 192, and synonymy.
Under stones, Home Lagoon, Hermite; July 1912; No. 106:
EONS
PILUMNUS CHRULESCENS A. Milne-Edwards, var.
? Pilumnus cerulescens Aleock, Jour. Asiat. Soc. Bengal, vol.
Ixvii. 1898, p. 196; Rathbun, Kgl. Danske Vidensk. Selsk.
Skrifter, 7 Rekke, naturv. og mathem., vol. v. p. 355, pl. 1.
fig. 15.
Under stones, Home Lagoon, Hermite; July 1912; No, 106:
orc
Dimensions. Length of carapace 12:6 mm., width 16°8 mm.
This is the same form as that which I recorded from the Gulf
of Siam (doc. cit.). The specimen is considerably larger than any
examined at that time, and some of the features are more sharply
marked. Although all of the regions are ornamented with
granules, they are not closely placed. The outer dentiform lobe
of the front is separated from the rest of the front by a U-shaped
gap. The two emarginations in the upper border of the orbit
are well marked. The antero-lateral projections and also the
subhepatic one are well developed spines; the orbital and sub-
hepatic spines are shorter than the other three; these last have
a stout base and form at the middle a shoulder from which a
cluster of hairs proceeds. A striking feature of the cheliped is
the presence on the upper margin of the merus of two strong
white spines, one terminal, the other subterminal and larger.
The carpus is armed with short, stout spines, the manus with
granules; neither spines nor granules are closely placed, and the
granules are absent from the lower distal corner of the larger
palm. The ornamentation of the whole upper surface is obscured
by the coarse hairs of uneven length.
The colour in formalin and glycerine is brownish; the carapace
has a ground of yellowish-brown, overlaid with patches of reddish-
brown ; upper surface of legs with two patches of brownish-red
on the merus, the carpus, and the propodus.
ACTUMNUS SETIFER (de Haan).
Actumnus setifer Alcock, Jour, Asiat. Soc. Bengal, vol. Ixvii.
1898, p. 202.
Monte Bello Islands: 1 ¢@.
The areole of the undenuded carapace are high and very deeply
separated and the lateral teeth very prominent; otherwise this
individual does not differ from typical specimens from Japan.
FROM THE MONTE BELLO ISLANDS, 661
Family OcyPoDID &.
Subfamily Ocypodinz.
Uca rorcipata (Adams & White). (PI. II. fig. 8.)
Gelasimus forcipatus Adams & White, Zool. * Samarang,’
Crust. 1848, p. 50.
3 d, large, medium, and small,
The cheliped of the largest specimen has the large distal tooth
of the dactylus at more than 4 the distance from the tip; in the
medium gpecimen, the tooth is at just the distal third. In the
smallest specimen this tooth is absent, the specimen representing
‘“‘form 2” of the species.
Subfamily Mictyrine.
Mictyris LoncicarPus Latreille.
Mictyris longicarpus Alcock, Jour. Asiat. Soc. Bengal, vol. bxix.
1900, p. 384, and synonymy.
Monte Bello Islands: 10 3,2 @.
Family [INACHID 2.
Subfamily Acanthonyching.
Hernia proteus (de Haan).
Huenia proteus Aleock, Jour. Asiat. Soc. Bengal, vol. lxiv. 1895,
p. 195, and synonymy.
Dredged off Hermite; July 12,1912; No.115: 19 immature.
Subfamily Pisinz.
Hyastrenus oryx A. Milne-Edwards.
Hyastenus oryx Alcock, Jour. Asiat. Soc. Bengal, vol. bxiv.
1895, p. 214, and synonymy.
Under stones in ‘lagoons’; June to Aug., 1912; No. 114:
OR uve
NAXIOIDES SERPULIFERA (Guérin), (PI. IT. figs. 9, 10.)
Pisa serpulifera Guérin, Icon. Régne Anim., Crust. pl. vii.
figs. 2, 2a, 2b, 2d. 1
Nawia serpulifera Milne-Edwards, Hist. Nat. Crust., vol. 1.
1834, p. 313.
Monte Bello Islands: 1 2 mature.
Under stones in ‘lagoons,’ June to Aug., 1912; No. 114: 1 9
Juv.
Dimensions. Length of carapace of large female to end of
horns, 92 mm. ; width, exclusive of spines, 59 mm.
An elongated sponge covers the upper surface of the right
662 MISS M. J. RATHBUN ON CRUSTACEA
rostral horn, and two other sponges are growing on the gastric
region.
The abdominal cavity is filled with young crabs in the adult
state. The cavity is about 40 mm. wide, 39 mm. Jong, and about
20 mm. high at the greatest extent ; the abdominal “appendages
are very slender, so that the bulk of the space is occupied by the
young, which number 162. These represent two stages, those of
the first or earlier stage being 13 in number with carapace about
3°5 mm. long; while those of the next or older stage are 149 in
number and about 5:°7 mm. long.
The younger ones although thin-shelled are harder and more
opaque than the next stage, and correspond to what is known
along the Atlantic coast of the United States as ‘ paper-shell ”
evabs ; ; they are covered with minute red pigment spots; the
carapace is almost smooth and naked, its shape is oblong, not
subtriangular as in adults; the postocular tooth is well-formed,
triangular and separated by a shallow sinus from the supraocular
eave; this latter shows no trace of a spine; as to the tip of the
rostrum, the inner of the two spines is well-developed and forms
the true end of the horn ; on its outer side there is a faint prom-
inence, which is later to become the strong lateral spine of the
adult; the eyes are large, protruding, the cornea of a light
br ownish-red colour.
By the next moult, which takes place within the brood-pouch
of the mother, the crab increases by more than half its former
size, and undergoes several notable changes. The carapace is of
similar form, bat the whole integument is soft and devoid of
colour spots; it is no longer smooth and naked, but uneven and
covered with crowded tubercles or granules, with the beginnings
of the more prominent tubercles of the ‘adult ; the surface is
more or less hairy, there being clusters of hooked hairs as in the
adult, and above all, a row on each rostral horn which is con-
tinued back on the carapace proper; the postocular cup forms a
tooth which is separated by a triangular sinus from the supra-
ocular eave, which last is armed witha small spine; rostral horns
elongate, each armed with two subequal spines. In the adult,
the postocular cup is separated from the supraocular eave only by
a closed fissure. This indicates that WV. serpulifera is generically
removed from the other Vaxioides and should be placed in the
neighbourhood of Lissa Leach (Zool. Mise. vol. ii. 1815, p. 69).
Subfamily Schizophrysine.
SCHIZOPHRYS DAMA (Herbst).
Cancer dama Herbst, Naturg. Krabben u. Krebse, vol. iil.
part 4, 1804, p. 5, pl. 59. fig. 5.
Schizophrys dama Miers, Challenger Rept. vol. xvii. part 49,
1886, p. 67. Alcock, Jour. Asiat. Soc. Bengal, vol. lxiv. 1895,
p- 245 and synonymy, but not “ Schizophrys aspera, p. pt.”
FROM THE MONTE BELLO ISLANDS. 663
Milne-Edwards, given in Kossmann’s synonymy; Illus. Zool.
‘Investigator,’ Crust. Part vi. 1898, pl. xxxv. figs. 2, 2a.
Under stones in ‘lagoons’; Monte Bello Islands; June to
Aug., 1912; No. 114: 1 2 juv. 20°3 mm. long, including rostrum,
14 mm. wide.
This specimen already shows the second or posterior spine on
the outer margin of the rostral horn; it is considerably smaller
than the anterior spine. It is not shown in the ‘ Investigator’
figure.
Distribution. Straits of Malacca (Alcock) ; West Australia,
3 to 5 fathoms (ers).
Family PARTHENOPID4S.
PartHENOPE (RHINOLAMBRUS) PELAGICA (Riippell).
Lambrus (Rhinolambrus) pelagicus Alcock, Jour. Asiat. Soc.
Bengal, vol. Ixiv. 1895, p. 267, and synonymy.
Home Lagoon, Hermite; June 13, 1912; No. 103a: 16.
Special locality not given: Go
Order STOM ATOPODA.
PRorosquInnA TRISPINOSA (Dana). (PI. IL. figs. 11, 12.)
See Borradaile, in Willey’s Zool. Results, Part iv. 1902, p. 400.
Dredged off Hermite, 3 fathoms ; in hole in piece of rock;
Silva, NOM pINow) Wi: Oe
Special locality not given: 1 @, variety.
Length of larger specimen (variety) 41 mm., of smaller speci-
men 25 mm.
The smaller specimen agrees with Borradaile’s amended
description and figure, except that the fourth abdominal segment
is corrugated much as the fifth is, only more faintly ; the sides of
the first, second, and third segments are also lightly carinated
with three or four smooth ridges.
The larger specimen, which I take to be the same species, has
some curious differences. The fourth abdominal segment and the
sides of the first, second, and third are nearly smooth, which
brings the specimen nearer to Borradaile’s, which was of the
same size. ‘he fifth segment is bordered posteriorly by fine
spinules. On the sixth and seventh segments, the spinules are
shorter and stouter than in the small specimen, telson considerably
wider than long, its three knobs less circular than in Borradaile’s
figure and in our small specimen, the outer knobs pear-shaped,
the inner one subtriangular; the marginal lobes are not curved
inward and are separated by shorter slits than in the smaller
specimen ; there is a carina parallel to the lateral margin, armed
like the margin with spinules.
Colour-markings. —Vhe specimens are preserved in formalin and
664 ON CRUSTACEA FROM THE MONTE BELLO ISLANDS.
glycerine. The larger one has small brownish spots regularly
arranged in transverse rows: two pairs on the carapace, one pair
anterior, one pair more widely separated, at the middle, and a
row of four on the posterior half; a row of four on the sixth
thoracic segment, and the first, third, and fourth abdominal seg-
ments; a row of two on the second, fifth, and seventh abdominal
segments. Knobs on telson olive-green, mottled. Uropods with
a broad band of reddish-brown across the middle. Swollen part
of chela white. The spotting of the smaller specimen is not so
distinct and there are in addition many finer, branched spots
down the middle of the animal, much as in Borradaile’s figure.
GoNoDACTYLUS CHIRAGRA (Fabricius) var. smirHit Pocock.
See Borradaile, in Willey’s Zool. Results, Part iv. 1902, p. 400.
Home Lagoon, Hermite, under rock; July 13,1912; No. 117a:
le he
‘his is the variety (Borradaile, loc. cit.), in which the keels of
the sixth abdominal segment and telson are considerably com-
pressed ; the keels of the sixth segment are produced without
constriction into long spines; the upper edge of the middle keel
of the telson is produced backward into a spine; and the flukes
of the anchor are formed by two narrow ridges running forward
from the hinder end of that keel. The dark spots on the first
five abdominal segments are not visible as the whole body is very
dark in the preserved specimen except the swollen part of the
chele, that of the manus being a deep blue, and of the dactylus a
pinkish-red.
EXPLANATION OF THE PLATES.
Prate I.
Fig. 1. Periclimenes hermitensis, type 2, side view, X 2.
. Periclimenes hermitensis, type 9 , rostrum, side view, X 5}.
. Periclimenes hermitensis, type 2, rostrum and antenne, dorsal view, X 44.
. Thalamita dispar, type 8, dorsal view, X 2.
. Actea glandifera, type 3, dorsal view, X 3.
Glyptoxanthus cymbifer, 2, dorsal view, X 2.
> OV 9 be
PuateE II.
Vig. 7. Glyptoxanthus cymbifer, type g, ventral view, * 2.
8. Uca forcipata, showing outer surface of chela of largest , X 11.
9. Nasxioides serpulifera, young, first stage, < 12.
10. Nawioides serpulifera, young, second stage, X 12.
11. Protosquilla trispinosa, 2, dorsal view, X 2.
12. Protesquitla trispinosa, variety, 2 , dorsal view, X 2.
ON MOLLUSCA FROM THE MONTE BELLO ISLANDS. 665
37. Report on Mollusca collected at the Monte Bello
Islands. By Tom Irepate *
[Received March 18, 1914: Read June 9, 1914. ]
(Text-figure 1.)
INDEX. Page
Geooraphicalerel ations Myspace seteee se eereeeeesescese-eeseicnitecinneaseie eters OOD
Bistrot Shellsicollectedteaee-.eetrernceecre costo ceserceeeeesesencaciaensccte crs: OOO
Systematic :— ;
J UETOR TACs, SIS 1D oocigeeesn sen abo pbnnOn ebb ox non DedccDoneEcHEEE | OL
Subularia montebelloensis, sp.N. ....... Be bon Cac coucer so Ree ere mn oH Ka)
Scaphella hedleyi, nom. nov., for 8. act depsmeeeacoaree od ON
The small collection brought back by Mr. P. D. Montague is
still of much interest on account of the geographical position of
the group.
Hedley, in the Proc. Linn. Soc. N.S.W. vol. xxvii. pp. 876—
883 (1903), introduced divisional names to indicate the different
faunal elements recognisable in the Marine Fauna of Australia.
He determined four primary divisions, to which he gave the
names Adelaidean, Peronian, Solanderian, and Dampierian
Regions. The limits of the Adelaidean Region were noted as
probably being Wilson’s Promontory in Victoria and Geraldton
in West Australia, embracing the whole southern coast of
Australia and round the south-west corner. The Peronian
Region designated the East Coast from Wilson’s Promontory
north to Moreton Bay in South Queensland. The Solanderian
Region was given to the remainder of the Queensland coast
northwards to Torres Straits; whilst the Dampierian Region
extended westwards from Torres Straits to Houtmann’s Abrolhos,
scarcely north of Geraldton, West Australia.
Verco (Trans. Roy. Soc. South Austr. vol. xxxvi. pp. 202-205,
1912) has now given a list of common Geraldton and Houtmann’s
Abrolhos marine molluses, which have been clearly shown to
belong to the Adelaidean Region, as out of a total of 150 species,
71 per cent. were also found in South Australia. This suggests
that the limit of the Adelaidean must be placed north of
Houtmann’s Abrolhos, and I can now show that few Adelaidean
forms extend as far north as the Monte Bello Group.
Hedley recently catalogued the Queensland Marine Molluscan
Fauna (Proc. Austr. Assoc. Adv. Sci. Brisbane, 1909, pp. 343—
371 & 809-810), when he mentioned over 1800 species. Very
numerous additions have since been made.
No list: of West Australian Marine Mollusca is known to me,
but it would be of great advantage to zoogeographers were such,
when prepared, shown under Hedley’s regional names.
* Communicated by Prof. J, Srantey Garprner, M.A., F.RS., F.Z.8.
666 MR. TOM IREDALE ON MOLLUSCA
The present collection only numbers forty-four species and
it is typically Dampierian; only one Adelaidean form, Conus
anemone Lamarck, occurring in it. The nomenclature and
sequence of species here used are based upon Hedley’s Queensland
List above noted, the few alterations made being accompanied by
notes which I submit to Mr. Hedley’s consideration. For it must
be acknowledged that all work on Australian molluses has
been rendered easy by the unparalleled energy displayed by
Mr. Hedley, and his knowledge of the literature and forms is
so complete that I know I am more likely than he to have
erred.
In the British Museum there is preserved a collection made at
the Monte Bello Islands and presented to that Institution by
Mr. T. H. Haynes. This collection has not been determined, and
includes about fifty additional species. I have not had time to
investigate the nomenclature of these forms, but as far as z0o-
eeosraphical relations are concerned they seem to confirm the
present collection in its entirety.
In the following List I note the word Solanderian against all
those included in Hedley’s Queensland List :—
Arca fusca Bruguiere, 1789, Solanderian.
Malleus malleus Linne, 1758, do.
Ostreea cucullata Born, 1778. do.
Chlamys radula Linné, 1758. do.
5 squamosus Gmelin, 1791, var.
55 lentiginosus Reeve, 1853, var, do.
Lima lima Linné, 1758, do.
,, multicostata Sowerby, 1843.
» fragilis Chemnitz.
Modiolus philippinarum Hanley, 1844. do.
Cardita incrassata Sowerby, 1825. do.
Cardium dupuchense Reeve, 1845. do.
is unedo Linneé, 1758. do.
Tridacna elongata Lamarck, 1819. do.
Antigona tiara Dillwyn, 1817. do.
Paphia titerata Linné, 1758. do.
Acanthopleura spinosa Bruguiére, 1792. do.
gemmata Blainville, 1825. do.
Haliotis sguamata Reeve, 1846. do.
= varia Linné, 1758, var. do.
Euchelus atratus Gmelin, 1791. do.
Turbo squamosus Gray, 1847, do.
Nerita albicilla Linné, 1758. do.
Acmeea saccharina Linné, 1758, var. do.
Cerithium fasciatum Bruguiére, 1792. do.
Hulima montagueana, sp.n.
Subularia montebelloensis, sp. n.
Cymatium aquatile Reeve, 1844. do.
Natica vitellus Linné, 1758. do.
FROM THE MONTE BELLO ISLANDS. 667
Cyprea caputserpentis inne, 1758. Solanderian.
5 caurica Linné, 1758. do.
x cylindrica Bon, ess do.
iS erosa Linné, 1758. do.
a errones Linné, 1758. do.
moneta Tiara 1758. do.
Scaphella volva Gmelin, Weeike do.
Bs zebra Leach, 1814, var. do.
a. hedleyi, nom. nov.
Voluta oblita Smith, 1909 (=norrisii auct.).
Ancilla elongata Gray, 1847. do.
Conus anemone Lamarck, 1810.
Arcularia suturalis Lamarck, 1822. do.
Rhodostoma auris-felis Bruguiére, 1789. do.
Bullaria columellaris Menke, 1843.
Note.—As regards generic denominations I would consider the
following alterations preferable :—
Cardium unedo should be Fragum unedo.
Ancilla elongata is Amalda elongata.
Arcularia suturalis , Alectrion suturalis.
The species noted seem to differ sufficiently from the types of
the genera first named to merit generic distinction.
Cardita incrassata Sowerby and Cyprea caputserpentis Linné
occur in Verco’s Geraldton List, but as both are marked as not
occurring in South Australia, they are obviously stragelers from
the north, and probably there reach their southern limit.
It will at once be observed that almost all the species occur
also in the Solanderian Region, whilst it is certain that some of
those not so marked do occur there, e.g. Lima multicostata
Sowerby*. The element characterising the Dampierian Region is
not well marked in the present eollection, the only notable species
being the Scaphelle, but it must again be observed that the
Adelaidean element is almost completely absent, which fact is of
some importance.
Mop10oLus PHILIPPrINARUM Hanley, 1&44.
Under this name I include specimens which I compared with
the presumptive type specimens in the British Museum. I, how-
ever, cannot see any differences worthy of consideration between
these and the type (presumably) specimens of J/odiola metcalfei
Hanley.
In the Proc. Zool. Soc. (Lond.) 1844, pp. 14-17, there is a
paper entitled “‘ Descriptions of a new species of Mytilacea &c.,’
by Sylvanus Hanley. This paper was read Feb. 13, 1844, and
published in July 1844. On p. 14 is described Modiola metealfei
from “ Hab. ?, Mus. Cuming, Hanley”; and on p. 15, JL. philippi-
narum from “Hab. Zebu, Philippinarum, Mus. Cuming, Hanley.”
* J note that Hedley (Mem. Austr. Mus. iv. 1902, p. 309) regarded this as simply a
variety of Lima lima Linné 1758, which would account for its omission from his
Queensland List.
Proc. Zoot. Soc.—1914, No. XLVi. 46
668 MR. TOM IREDALE ON MOLLUSCA
If this were the first introduction, I/. metcalfei has priority, but
in Hanley’s ‘ Bivalve Shells,’ these two species are again described
and figured. ‘The locality given for both species is the Philippine
Tslands, and the specimens above noted from the Cuming Collection
are so labelled. Both names appear on the same page, but here
again I. metcalfei appears first. According to the data given
in the Introduction to the work this page appeared in 1843!
Whichever appeared first, I conclude that MZ. metcalfer would
claim usage in preference to MZ. philippinarum. Ii 1t be conceded
that both species were described from the same locality, there
cannot be the slightest hesitation in accepting the identity of
the two species, the very slight difference in form being almost
certainly due to age.
ANTIGONA TIARA (Dillwyn, 1817) *.
In illegally rejecting Cytherea of Bolten 1798 and accepting
Antigona Schumacher 1817 in its stead, Jukes-Browne (Proce,
Malac. Soc. (Lond.) vol. xi. p. 70, 1914) has unwittingly selected
the most appropriate name. For a valid reason for the rejection
of Cytherea Bolten 1798 exists in the fact that there is a prior
Cytherea of Fabricius, Ent. Syst. vol. iv. p. 413, 1794, as well as
a Cythere O. F. Miller, Entomostraca, p. 63, 1785. This was
unknown to Jukes-Browne and overlooked by Dall, Hedley,
and Suter, who have recently used Bolten’s name. According to
nomenclators, Antigona Schumacher 1817 was predated by Aznta-
gonus Htibner 1816, and hence, according to British usage,
invalid ; but Sherborn has shown that Antigonus was not
published by Hiibner until 1820, leaving Antigona unassailable.
This detail was also unknown to Jukes-Browne as to most
malacologists.
ACANTHOPLEURA SPINOSA (Bruguiere, 1792).
Specimens of large size were obtained, and these seem inter-
esting on account of the southern distribution, this being the
furthest south record I have traced.
ACANTHOPLEURA GEMMATA (Blainville, 1825).
For the past five years I have been studying the forms of the
genus Acanthopleura, especially with relation to those grouped by
Pilsbry (Man. Conch. vol. xiv. pp. 221-226, 1893) under the name
Acanthopleura spiniger Sowerby. This would seem an appropriate
place to note generally the points raised.
Hedley has accepted for the species name Linné’s Chiton
aculeatus, but I agree with Pilsbry in rejecting this as indeter-
minable. Pilsbry, however, discarded Chiton gemmatus Blainville
(Dict. Sci. Nat. vol. xxxvi. p. 544, 1825) and selected Chiton
spiniger Sowerby (Mag. Nat. Hist. 1840, p. 287, Suppl. pl. xvi.
fig. 2) instead.
* {The parentheses around the names of authors placed after scientific names in
this paper are used in accordance with Article 23 of the International Rules of
Nomenclature (Proc. 7th Int. Cong. Boston, 1907, p. 44 (1912))—Epiror. |
FROM THE MONTE BELLO ISLANDS. 669
T can see no flaw in Blainville’s description, and the locality
“New Holland” suggests that it might have been brought home
from Torres Straits where it was very common, or it might even
have been collected by Peron and Lesueur at Shark’s Bay, West
Australia, whence Thiele (Die Fauna Siidwest Australiens,
vol. i. p. 398, 1911) has recently recorded it under the name
Acanthopleura spuriger Sowerby.
Sowerby’s Chiton spiniger was of unknown locality, and the
figure 1s somewhat abnormal as to the length of the spines on
the girdle.
Pilsbry included A. spiniger Sowerby in the typical subgenus
(A. spinosa (Bruguiere) being the type of the genus) and proposed
a new subgenus Amphitomura (Nautilus, Jan. 1893, p. 105) for
Ch. borbonicus Deshayes, admitting Ch. brevispinosus Sowerby
as distinct from that species but referable to the same subgenus.
These two are practically from the same locality, and typical
specimens prove their identity.
From the Red Sea comes a form which has just as commonly
been referred to A. “ spiniger” as to any other species, Pilsbry
making note of this. I have examined ma ny Specimens, and this is
undoubtedly referable to the species brevispinosa, but as certainly
subspecifically separable. This form, which should bear Roche-
brune’s name balanse, completely breaks down any subgeneric
distinction between brevispinosa and gemmata; but the latter is
just as clearly subgenerically recognisable when contrasted with
spinosa.
Pilsbry included Ch. echinatus Barnes under Acanthopleura,
though forming a subgenus (Jesotomura, Nautilus, Jan. 1893,
p. 103) for it. I would reject this species from the genus, so that
my genus Acanthopleura would read :—
ACANTHOPLEURA.
Subgenus Acanthopleura.
spinosa Bruguiere.
Subgenus dAmphitomura.
brevispinosa Sowerby
(=borbonica Desh.)
with several subspecies.
gemmata Blainville
=spuriger Sowerby)
with several ‘subspecies.
Subgenus Maugeria.
granulata Gmelin.
I must note that dissection of the type of Chiton cunninghami
Reeve, described from ‘ Australia,” proves that shell to be
identical with brevispinosa and the locality incorrect.
TURBO SQUAMOSUS Gray, 1847.
Hedley included in his List a Turbo foliaceus Philippi, 1846.
In the Zeitschr. fiir Malak. (Menke) 1846, p. 98, Philippi
46*
670 MR. TOM TREDALE ON MOLLUSCA
described Turbo lamellosus. In the Conch.-Cab. {Kiister) Turbo,
p- 41, Philippi figured this species, but renamed it Turbo foliaceus
as his former name was preoccupied by Broderip. This would
seem to be the entry quoted by Hedley, the titlepage of the
volume giving 1846. In the British Museum copy of this work
the dates of publication have been collated, as the titlepage date:
veferred only to the first few pages. I there find that page 41
appeared in 1847.
Gray, in the Narr. Surv. Voy. ‘ Fly,’ vol. ii. p. 359, fig. 8, pl. 1.
1847, described the same shell from Port Essington under the
name J'urbo squamosus. There is no question of priority, how-
ever, as there is a prior J'urbo foliaceus Gmelin (Syst. Nat. p. 3602,
1791) invalidating Philippi’s name, :
I have noted that Reeve (Conch. Icon. Turbo, fig. 17, 1848)
gave the name 7urbo laminiferus to the same species, and that
specimens from Torres Straits were independently named Z'urbo
foliaceus by Hombron and Jacquinot, which name was published
by Rousseau in the Voy. Péle Sud, vol. v. 1854, p. 60. The
ficures (Moll. pl. xiv. figs. 34-37) may have been issued earlier,
but I have no data, and the invalidity of the name obviates any
inquiry.
AcM#A SACCHARINA (Linné, 1758), var.
Under the above name I have included two specimens.
In his Queensland List Hedley admitted two species of Aemea
under the names, Aemcea costata Sowerby 1839, and dAemaa
saccharina Linné 1758. Why the former name was used I cannot
say, as in the Proc. Linn. Soc. N.S.W. 1904, p. 189, Hedley himself
went into the matter of the nomenclature of the Sydney shell
known as Acmca costata Sowerby and endeavoured to prove that
name inapplicable, and that the correct name was dAemea alti-
costata Angas (Proe Zool. Soc. (Lond.) 1865, p. 56, pl. ii. fig. 1)
given toa South Australian form. I have not seen any contro-
version of Hedley’s argument, so with the help of Mr. G. C,
Robson, of the British Museum, I tried to clear the matter
up. Working at this group Mr. Robson recovered the spe-
cimen from which the figure of Lottia? costata Sowerby (Zool.
Beechey’s Voy. 1839, p. 147, pl. xxxix. fig. 1) was prepared, and
it proves to have the data “ Arica, Peru” on the back of the
tablet. It is obviously not the Australian shell, and this discovery
absolutely disposes of Sowerby’s name as referable to the Sydney
Acmea.
The same species occurs in South Queensland, but in North
Queensland it is replaced by a different species, which Hedley
catalogued as Acmea saccharina Linné 1758. At Port Curtis.
I collected a series of specimens, and the determination of the
name to be used has caused quite a lot of trouble.
Mr. Robson has investigated the question of the type locality
of Linné’s Acmea saccharina and has fixed this as the Philippine
FROM THE MONTE BELLO ISLANDS. 671
Islands. This is necessary, as there can be no question that, due
to the wide range of the species, it can and must be divided into
subspecies. In the latest Monograph of the family, Pilsbry (Man.
Conch. vol. xiii. p. 49 et seg., 1891) did not determine these but
admitted demca saccharina Linné, quoting as a synonym Patella
lane Reeve: p. 50, var. stellaris Q. & G., giving as synonyms
stella Lesson and %octoradiata Hutton, and naming p. 50, var.
perplexa nov., from Australia, quoting under this name 2 stellaris
Reeye and octoradiata Hutton. He accepted Sowerby’s name
costata for the Australian shell, synonymising with it Angas’s
alticostata. In the Proc. Linn. Soc. N.S.W. 1904, p. 188,
Hedley put on record that Pilsbry’s var. perplexa was absolutely
Hutton’s octoradiata, and that this was a valid species. There
can be no question about this, but Hedley accepted Hutton’s
name, whereas he should have chosen Pilsbry’s, as Hutton’s name
was invalidated by Patella octoradiata Gmelin, Syst. Nat. p. 3699,
1791.
Suter, reviewing New Zealand Acmeide (Proc. Malac. Soc.
(Lond.) vol. vii. pp. 315-326, 1907), recognised in Patella stella
Lesson (Voy. ‘Coquille,’ vol. 11. p. 421, 1831) the shell commonly
known as corticata Hutton, but admitted the form Hutton named,
with subspecific rank. He also pointed out its alliance with the
Australian 4. alticostata Angas. The New Zealand species is
- certainly specifically distinct from every form of sacchurina.
At the same place Suter admitted octoradiata Hutton, and
placed these two species in a subgenus Collisellina. This name
was proposed by Dall (Amer. Journ. Conch. vol. vi. p. 259, 1871)
for his section B, which covered saccharina Linné and borneensis
Reeve. In the Voy. de l’Astrol., Zool. vol. iii. pt. ii. p. 349, 1835,
Quoy and Gaimard introduced their genus Patellotda for southern
Limpet-like forms with distinct anatomical features. In 1847
Gray designated as type P. rugosa Q. & G., and as this is one of
the original members of the genus this selection must stand.
I suggest that P. rugosa Q. & G. is a member of the present
group, and that Patellotda Quoy and Gaimard should displace
Collisellina Dall. The group is well defined and easily recog-
nisable.
Patelloida stellaris Q. & G. for a New Ireland shell, which is
certainly not identical with the North Australian form, is invalid
on account of Bolten’s Patella stellaris (Mus. Bolten. p. 12, 1798)
given to a different form of A. saccharina.
Patella lanw Reeve was described from Japan.
Other early names given to saccharina-like shells are all in-
applicable to the Australian form, which is therefore unnamed.
These differ appreciably from typical specimens, whether specimens
from Monte Bello Islands, Port Essington, Cape York, or Port
Curtis are contrasted, and I therefore note them as 4. saccharina
Linné, var.
Verco referred the shells from South-West Australia to 4. alti-
costata, and specimens in the British Museum labelled West
672 MR. TOM IREDALE ON MOLLUSCA
Australia and Swan River, West Australia, confirm this dis-
position, as they closely approach the South-Kast Australian species
and are very different from any form of Aemea saccharina.
These sketchy remarks must well show the confusion that exists
in this family, and the urgent need of ‘a skilful and careful
monographer.
My notes would read easily if summarised thus :—dAemea
saccharina Linné. ‘Type locality, Philippine Islands, divisible
into subspecies: Patella lanx Reeve, Japan, one valid name.
Patelloida stellaris Quoy & Gaimard, New Ireland: form probably
recognisable, name invalid, Subspecies ranging from Cape York
westwards to Monte Bello Islands and southwards to Port
Curtis, recognisable but unnamed.
Acme saccharina, var. perplexa Pilsbry is a distinet species,
commonly called Aemea octoradiata Hutton, but Hutton’s name
is Invalid.
Acnuea alticostata Angas is the name for the Sydney shell,
which ranges southwards through Bass Straits and then west-
wards to Geraldton, West Australia, and this is quite a valid
species. Sowerby’s Lottia ? costata (type preserved in the British
Museum) came from Arica, Peru, and has no connection with the
Australian shell erroneously so-called,
Acmea stella Lesson, from New Zealand, is closely allied to
A. alticostata Angas, bar is certainly separable as a distinet
species.
The group is well marked and has been classed under Colli-
sellina Dall 1871, but Quoy and Gaimard’s genus name Patelloida
seems to claim usage on account of Gray’s designation of P rugosa
Q. & G. as type. I regard this species as certainly referable to
this group.
EULIMA MONTAGUEANA, sp. n. (Text-fig. 1 A, B.)
Shell of medium size for the genus, thin, smooth, solid, glassy,
imperforate, not translucent, variced, many-whorled, sutures
impressed. Colour milk-white. In shape it 1s she anply conical
with the spire somewhat tending backwards. The largest specimen
has the apex missing, but fourteen whorls remain. The next in
size, which I select as the type, has the apical whorl somewhat
bulbous and succeeded by fifteen whorls, the basal three or four
whorls showing a peripheral keel. Varices regularly sueceeding
and advancing | spirally can be observed on the last ten whorls -
on the spire shins these are too indistinct for recognition.
Aperture obliquely pyriform, outer lip simple but not thin,
base somewhat contracted; columella straight and reflected as a
slight callus which extends across the body-whorl to join ‘the
outer lip at the posterior angle.
Length of type 17-5 mm., “breadth 5:5 mm.
Pavia. Off Hermite Island, Monte Bello Group. Dredged in
4 fathoms
FROM THE MONTE BELLO ISLANDS. 673
Remarks. Melanella Bowdich 1822 has been shown to have
priority over Hulima Risso 1826; but as the type of the former
is a “humpbacked” species it has been suggested that these
might be separated generically, and Hulima might be retained
with the conventional usage.
I have not yet examined the type of Melanella, but Austra-
lasian “ humpbacks ” seem to have apical features very distinct
from species of Hulima occurring with them.
Text-figure 1.
’
Cc
A, Side view of Hulima montagueana showing lateral ascending varices.
B. Front view of Hulima montaqueana.
C. Subularia montebelloensis.
SUBULARIA MONTEBELLOENSIS, sp. n. (Text-figure | C.)
Shell small, thin, not translucent, slender, aciculate, imper-
forate, glossy, white, many-whorled, smooth. Apex very slender,
the apical whorl minute with sixteen succeeding whorls, the
sutures indistinct but followed by a transparent band, the
remainder of the whorl milky.
The last whorl has the periphery rounded and varices do not
appear to be present. The aperture is oval, anteriorly broadened
and subehannelled ; columella truncate, advancing obliquely and
slightly reflected as a callus extending across the body-whorl,
Length of type 12°5 mm., breadth 3 mm.
674 MR. TOM IREDALE ON MOLLUSCA
Habitat. Off Hermite Island, Monte Bello Group. Dredged in
4 fathoms.
Remarks. A vague resemblance to Lulima acicula Gould may
account for the previous non-recognition of this species. Tryon
(Man. Conch. vol. viii. 1886) synonymised with that species,
aciculata Sowb.. pyramidalis Sowb., and vitrea A. Ad., an in-
congruous combination needing no criticism, but since that time
little work appears to have been done on this group as regards
Indo-Pacific species.
Tryon placed these in Zulima in the section Subularia Monte-
rosato, which name was proposed to replace Leiostraca Adams,
which was considered preoccupied. As there is a prior Leio-
stracus as well as Liostraca and Liostracus, | would agree in
rejecting Letostraca.
This species is not a typical Subularia, but might be refer-
able to Monterosato’s section Acicularia, but that name 1s pre-
occupied.
Natica virecivs (Linné, 1758).
I had determined the Monte Bello shell as Verita rufa Born,
and noted its absence from Hedley’s Queensland List, before I
received my friend’s latest paper (Proc. Linn. Soc. N.S.W.
vol. xxxvili. 1913), where on p. 299 he recorded this species from
Palm Island, Queensland, and noted that the correct name for
the species was as given above.
SCAPHELLA ZEBRA Leach, 1814, var.
The shells do not agree with typical S. zebra, from the East
Coast of Australia. In the British Museum, similar shells from
West Australia were named S. ellioti Sowerby. Upon investi-
gation this name was found to have been given to a North
Avstralian shell (Journ. de Conch. 3rd ser. vol. iv. p. 358, 1864;
vol. v. p. 25, pl. 11. fig. 19, 1865), and the figure did not agree
with West Australian examples. As far as I can trace, no name
has yet been given to the West Australian shell. I, however,
forbear its nomination, and in preference record it as above:
it cannot be called S. tuwrneri, as the figure of Voluta turnert
Griffith and Pidgeon (Anim. Kingdom Cuy, vol. xii. Moll. pl. x.
fig. 1, Index, p. 601, 1834: no locality given) shows quite a
different shell.
SCAPHELLA HEDLEYI, nom. nov.
Scaphella reticulata (Reeve) does not appear in Hedley’s
Queensland List, but was added to that fauna by Shirley (Proce.
Roy. Soc. Queensland, vol. xxiii. p. 99,1911) from the Gulf of
Carpentaria.
The species was described by Reeve in the Proc. Zool. Soe.
(Lond.) 1845, p. 144, under the name Voluta reticulata, and
figured in the Conch. Icon., Voluta, sp. 25, pl. xi. figs. 25, a-b.
FROM THE MONTE BELLO ISLANDS. 675
That name, however, had been previously utilised by Linné
(Syst. Nat. 12th ed. p. 1190, 1767) and Martyn (Univ. Conch.
vol. iv. fig. 126, 1787), and as I have noted no synonymy I
propose to rename Reeve’s species as above. It gives me great
pleasure to associate with such a beautiful shell the name of my
friend Mr. C. Hedley. Sowerby’s V. gatliffi, described from North
Australia, differs in shape as well as coloration, and I know no
other species that can be compared with this.
Conus ANEMONE Lamarck, 1810.
Tn his latest paper cited above Hedley (p. 307) gave some notes
on this species, observing that the West Australian shell seemed to
be typical and noted it from Port Essington and also Tasmania.
My specimens from Monte Bello Islands agree with his conclusions,
so I anticipate that Verco’s Geraldton record would also agree. If
it be conceded that the East Australian shells are only sub-
specifically distinct (my own shells from Sydney, New South Wales,
and Lord Howe Island agree with Hedleys remarks) then the
names to be used would be
Conus anemone Lamarck. North, West, and South Australia ;
of which C. novehollandie A. Adams is an absolute synonym ;
and
Conus maculosus Sowerby. Hast Australia and Lord Howe
Island ;
with jukesii Reeve, maculatus Sowerby, and rossitert Brazier, as
absolute synonyms.
R#HODOSTOMA AURIS-FELIS (Bruguiére, 1789).
The genus name here used was introduced by Swainson, Treat.
Malac. pp. 208, 344, 1840, the species cited being coffe Chemn.
120, f. 1043; fabula Feér. Tab. Syst. 105, n. 24; and nucleus id.
ib. n. 26. In the Tab. Syst. Moll. Férussac, p. 109 (or 105)
included fabula in Auricula (Conovulus), and then proposed
Auricula (Cassidula) felis Lamarck=coffea Chemn., and added
Auricula nucleus without any subgeneric designation. Cassidula
has been used for the group typified by auris-felis Bruguiére
(=coffea Chemn.) ever since this introduction, which only dates
from 1821. In the Syst. Anim. p. 348, 1801, twenty years pre-
viously, Lamarck had proposed Cassidulus for an Echinid, which
name is now generally accepted as invalidating Cassidwla.
ON CEPHALOPODA FROM THE MONTE BELLO ISLANDS. 677
38. Cephalopoda from the Monte Bello Islands.
By G. C. Rozson, B.A.*
[Received March 18, 1914: Read June 9, 1914. |
(Text-figure 1.)
INDEX.
Structure : Page
Radula, mandibles, poe ee of Sepiadariwm
CUFURHT Oy SDo1s- sonaccocooc9 sosonayocconscoodeacceannne LOE (ots)
Variation, &c.:
Differences between Peay eee and Sa ees
stunmarized . paene ee 677
eeuie sar imm natnibuam, sp. n., Gntern me vediate betwe een
epiadarium and Sepioloided .............20000-..... 877
Geographical Zoology :
Distribution of Sepiadariwn auritum, probably
DUO MALO USA eae ee ee ee Ons
Systematic :
SCPC PONTO. GOUFUCRUID, SOs Ne cosooenceacdonscosespouane Whe’
Of the two species of Cephalopoda obtained by Mr. P. D.
Montague from the Monte Bello Islands, one specimen is of
considerable interest—-a Myopsid which has been referred after
some hesitation to Sepiadarium.
The differences between the latter genus and the closely-allied
Sepioloidea, though they have been touched upon by several
authors, have not been fully summarized as yet. The following
table gives the more important differences :—
Sepradarium Steenstrup. Sepioloidea d’Orb.
1. Funnel attached to mantle by a .... by cartilaginous plugs of the
ligament (1) (2). mantle that fit into sockets on
the funnel (1) (2).
2. Ventral pores absent (2). .... present (2).
3. Mantle fringe absent (2). .... present (2).
4. Fins short (4). 65 0 komme (2);
5. The hectocotylized arm bearing Suckers of the hectocotylus per-
on its distal half a series of sisting as small papille ; the arm
transverse ridges, which are the grooved diagonally on its inner
laterally produced peduncles of side (4).
the suckers (4).
In respect of Nos. 1, 2, 3, and 5 in the above list, the form
here described is referable to Sepiadarium. The length of the
fins, on the other hand, suggests aftinity with Sepioloidea. It is
not desirable on such slender grounds, however, to create a new
genus intermediate between the two forms under discussion for
the reception of this species. But it is certain that in the
present state of our knowledge we are entitled to regard this as
an unusual form of Sepiadariwm, intermediate in respect of one
character between that genus and Sepoloidea.
* Communicated by Prof. J. Sranzey Garprner, M.A., F.RS., F.Z.8., and
published by permission of the Trustees of the British Museum.
678 MR. G. C. ROBSON ON CEPHALOPODA
Up to the present Sepioloidea appears to be regarded as the
Australian form of the two Sepiadarian genera, while Sepia-
darium is considered the Pacific form. We now find that the
latter extends its range to W. Australia, though not by a typical
member of the genus. Whether, in the first instance, the
distinction of the two forms into Pacific and Australian rested
upon secure and sufficient evidence we camnot say as yet. If the
investigation of the Cephalopod faunas of these areas upholds
this distinction, the interest and importance of this species of
Sepiadarium with Sepioloidea-chavacters, as occurring in the
distributional area of the other genus, will be increased.
The structure of the hectocotylus corresponds closely with that
described by Brock (5) for Sepiadarium. It should be observed
that the series of transverse ridges or bars (Brock’s ‘‘ Quer-
balken”) are of such a shape as to suggest the obvious con-
clusion that they represent the fused bases of the pairs of
suckers, a proximal member of the morphologically posterior
row being fused with a distal member of the morphologiealiy
anterior row.
1. SEPIADARIUM AURITUM, sp. n. (Text-fig. 1.)
External appearance.—The animal is small and squat, the
width of the mantle area being about equal to its length. The
fins are rather long and ear-shaped, the inferior portion being
slightly broader than the superior. ‘The mantle-insertion at the
neck is tolerably broad, while posteriorly the edge of the mantle
exhibits a very slight concavity.
There is a very feebly developed interbrachial membrane.
On the arms the suckers are arranged in two alternating rows,
save at the distal third, where they become abruptly smaller and
more irregularly disposed.
The tentacular arms are provided with a short membrane
conterminous with the area occupied by the suckers.
The colour (formalin-preserved specimen) is dull grey, covered
on the dorsal surface by numerous small round patches of pale
red or brown and fine black or dark-brown spots. Only the
latter are continued on to the ventral surface and the arms. The
patches and spots are found, though more sparsely, upon the fins.
For character of the hectocotylized arm, v. text-fig. 1, C.
Dimensions :—
Mantle, max. length 00. 11:25 mm.
- max. breadth’ -9°%).\' 0. ea ae
Wencthror tinsm tees: wn toe. 8°75
Total length (from apex of mantle
to interbrachial membrane of
the weulbia) aeMis)\ sg... a 17
Length of arms: Ist pair......... 7
PASC Lape aes es Se 6°8
9
7
FROM THE MONTE BELLO ISLANDS. 679
Text-figure 1.
—
cy He Mb :
|
Sepiadarium auritwm.
A. Dorsal view. X2: a, outline of fin.
1. Attachment of funnel to mantle. X 2.
C. Hectocotylus. x 6.
D. Mandibles. x 12.
E. Radula. 4 oc. X 6 obj.
680 ON CEPHALOPODA FROM THE MONTE BELLO ISLANDS.
Internal characters.—The mandibles (text-fig. 1, D).
The radula (text-fig. 1, E) resembles that of S. kochii figured
by Appellof (3) pretty closely ; but differs in the condition of the
basal plate of the median tooth, which is much deeper and of a
different shape, in certain characters of the lateral tooth and of
the inferior marginal. It is unfortunately impossible to give a
fuller account of the internal characters, owing to the fact that
only one specimen is available for examination.
Locality. Dredged off Hermite I. (Monte Bello Islands),
W. Australia.
Type in the British Museum (Zoological Department).
2. POLYPUS sp.
Two specimens from the same station (one immature).
This form, which might be referable to more than one Pacific
species, cannot be satisfactorily identified.
Literature referred to.
. JouBIN, L. Meém. Soc. Zool. France, xv. 1902, p. 81.
. Sreenstrur, J. K. Danske Vidensk. Selsk. Skrift., 6 Raekke,
1881, p. 214.
. Appettor, A. Abh. Senckenb. Naturf. Ges., Bd. xxiv. 1898,
p- 061.
Petseneer, P, Lankester’s Treatise of Zoology. Mollusca.
Brock, J. Zeitschrift fur wiss. Zool. xl. 1884, p. 105,
we
ae
ON A NEW LIZARD FROM THE CANARY ISLANDS. 681
39. Description of a new Lizard from the Canary Islands *.
By Dr. Pu. Lenrs f.
[Received May 14, 1914; Read May 19, 1914. |
LACERTA CHSARIS.
Abstract P. Z.8. 1914, p. 41 (May 26th).
Physiognomy and general proportions of Lacerta galloti, but
much smaller and, from a phyletic point of view, of a more
primitive pattern of coloration.
Head rather large, difference in size according to sexes not so
marked as in Z. galloti and its larger allies. Length of head
1 of length to vent in the male, very little less in the female.
‘he cheeks not noticeably swollen in the male. Width of head
+ of the length, depth 3, width of pileus 4 in both sexes. Snout
as long as the postocular part of head (to posterior border of
tympanum) in the female, slightly longer in the male.
Weck distinctly narrowed. ody slightly flattened.
Tail long, more than two-thirds total length.
Limbs elongate, the fore limb im both sexes reaching the
nostril, the hind limb reaching the collar or not quite so far in
the female, a little beyond in the male.
Rostral usually touching the nostril {, separated from the
frontonasal.
Frontonasal as long as broad or slightly longer.
Frontal a little shorter than its distance from the end of the
snout in the female, but slightly in the male; twice as long as
broad (on its narrowest point) in the female, slightly longer in
the male; not touching the first supraocular.
Supraoculars separated from superciliaries by a series of
granules.
Parietals as long as, or slightly longer, rarely shorter § than
the frontal in the male, usually a little longer in the female,
strongly bent down on the temple, occupying the place usually
held by the supratemporals ||.
Occipital at least as broad as the interparietal, usually con-
siderably broader, and constantly broader than long; on an
average larger in the male than in the female 4.
Asingle postnasal. 5 upper labials anterior to the subocular **.
* [The complete account of the new species described in this communication
appears here, but since the name and a preliminary diagnosis were published in the
‘Abstract,’ No. 134, 1914, the species is distinguished by being underlined.-—
Eprrox, |
+ Communicated by Dr. G. A. BounenGErR, F.R.S.. F.Z.S.
{ Exceptions rare: for instance in Nos. 3 and 14 of the annexed table.
§ Fer instance in No. 13.
|| The author does not wish to support the theory of a fusion of the said shields.
As is generally the rule in the species of Lacerta.
abe superposed anterior loreals on each side in No. 4, double anterior subocwars
in No. 12.
DR. PH. LEHRS ON A NEW LIZARD
aie SE
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rs Gh. Ep ca Ss Se eS to vent.
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xn Ch Se SS a oo :
ie EE Me ey oT el eee Lane ars Width and length
a Pepe ee Be ob Pw eH wp SD of occipital shield.
bo Or bo or “I “I ~T bo bf bo
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2 m i © GG 9 MSs iss 8) 1 :
o bt oo CO oa coe (a) (S) = oO = Gular scales.
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bo to eg © W fs OS)
® oo tm tw = & S' S fs 2 & Femoral pores
= LS ESS eS! 0S) TS Rey Se RS es r] i
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Sonata
2 Y Con TSE Gog CON eke Loe Subdigital lamelle.
FROM THE CANARY ISLANDS, 683
Temporal scutellation granular, often finely, with more or less
developed masseteric and tympanic shields; a series of small
supr atemporals.
Scales covering the lower eyelid extremely small and granular,
those in the centre scarcely, if at all, differentiated.
Dorsal scales very small, rhombic- granular, more or less dis-
tinctly keeled, 90-110 across the middle of the body; 3 or 4
series corresponding to one ventral plate.
Upper caudal scales much smaller than the dorsals.
Gular scales 33-44 on a line between the collar and the third
pair of chin-shields; gular fold distinct, collar not serrated,
composed of 7-11 plates.
Ventrals in 10 or 12 longitudinal and 29-31 Ponsvene series.
Anal comparatively small.
Femoral pores 24-31.
Subdigital lamelle 30-36 under the fourth toe.
Coloration (from life). Pileus of a dark or light bronze-brown,
more or less profusely speckled with black.
Superciliary streak, if distinct, on each side running entirely
upon the parietal and continued to the base of the tail, or
beyond; this streak of a bright yellow in life.
A light streak along the spine, well defined from the bronze
colour of the dorsal region.
A short subocular streak, of a duller yellow than the super-
ciliary, and often interrupted, extends a little way beyond the
tympanum in the middle of the dark brown temporal band,
which is edged below by a continuous dull yellow streak, extend-
ing from below the tympanum to the base of the hind limb,
reappearing again on the side of the tail.
The dark bands in some specimens (young males) with small
light, dark-edged spots without any tendency to form transverse
series. Larger bright spots on the limbs.
Lower surface of head blackish grey, or even black in full-
grown specimens; the dark shade sometimes extending to the
breast. Belly dirty white, or pale greyish, without any spots ;
a few roundish, ileal white spots sometimes present on the
margin.
Frequently a small sky-blue spot on the dark temporal band
above the insertion of the fore limb, another of the same colour
on the upper arm. In a few adult females the whole lower
surfaces of the thighs of the same bright blue.
In some fully adult males the markings become very indistinct
or may nearly entirely vanish, such specimens being blackish
brown.
Iris silvery.
Habitat: Hierro, Canary Islands; a large number of speci-
mens were collected by Dr. Caesar Boettger at Las Lapas, in
the “ Golfo” of Hierro, and presented to me in August, 1913.
The types, which I received alive, are preserved in my private
collection. é
This species belongs to the same group as the well known
Proc. Zoou. Soc. 914, No. XLVII. 47
684 ON A NEW LIZARD FROM THE CANARY ISLANDS.
L. galloti D. B., from which it differs to the same extent as
LL. simonyi Steind., from the “ Roques del Zalmore,” near Hierro,
but in an opposite direction.
' It differs from JZ. galloti in its smaller size, in its broader
occipital shield, in the downward extension of the parietal shields,
in fewer longitudinal rows of ventral shields (10-12, instead of
12-14), in the higher number of dorsal scales (90-110, instead of
80-98), in the more primitive pattern of coloration, as well as in
the feeble degree of secondary sexual differentiation.
In my opinion it represents the most primitive type of the
L. galloti group.
ON THE MECHANISM OF SUCTION IN LYGUS. PABULINUS. 685
40, The Mechanism of Suction in the Potato Capsid Bug,
Lygus pabulinus Linn. By P.R. Awart, B.A. (Cantab.),
D.LC.(Lond.), Sir John Wolfe-Barry Research Scholar,
Imperial College of Science, London *.
[ Received January 31, 1914: Read March 17, 1914}
(Text-figures 1-29.)
INDEX. Page
Introduction ..... BA ro ACS DNE ERE HUST IO EUnet RBS Geos ape ETE oF =D
Material and Method .. PREP Ra Meatainngein en ptt ee eae) ORG.
Hiomolerencothenviont erat Bopean esa cua cunbondtos68 doc aastas Doon oleH7/
Moxpholosyotstheviea dm rieece nesters cetseeenchcckc emcees ore cern Oe
Tentorium ..... sel Bua edauels cutaiscon sate Aveueaeas son Mer Meaty ia AO
Muscles of the Head RA ate dean can enero e cae ita) APD
LEVEN a1ixg 8 For Hse a Ghanseidoeusacoa Da adelaaeetie ee On terlace Cac Roe a see pene eee aA LS)
Gustatory Cues Sot nice cada care Dap caso BA caamec Doaaeouee EE MCE ERY OT
PUMIp-ap Parabus ceawes see eee ese tee hs censotem ee encca teens teres CO
Mechanism pacncnion Bn Renee Suen ee nantaNy eR Re a TA eae aR ees LO
B31) StOyeRIEY OD) Fae ena neaasuees anu coc aBeNs ode Gee Hes ectdunas ScbaC ee noeseRe omen 732
INTRODUCTION.
The investigation of the mechanism of suction in Lygus
pabulinus arose out of an inquiry by Professor Lefroy, of the
Imperial College of Science, and Mr. Horne into the production
by insects, on the foliage of the potato, of symptoms resembling
those produced by fungi and by bacteria. They had produced
definite symptoms by the infection of potato foliage with a Capsid
(Lygus), a Jassid (Hupteryx), and other sucking insects; the
symptoms were markedly characteristic of the differ ent species ;
and the investigation of the actual mechanism of suction thus
became necessary. I took up this inquiry at the suggestion of
Professor Lefroy, who was in India at the time.
The insect is common on the potato and other plants during
the summer. More than one species of Lygus is concerned with
the damage to the plants. The species used in this investigation
has been identified by Mr. W. L. Distant at the Natural History
Museum as Lygus pabulinus Linn.
The scope of this paper is limited. It does not pretend to
give full anatomical or morphological descriptions, but it describes
the different structures of the head, both morphologically and
anatomically, as far as they appear to be important in the
mechanism of suction.
I very gratefully acknowledge my indebtedness to Prof. Lefroy
for his help and encouragement during the whole of my work, to
Mr. Clifford Dobell for his valuable criticism and suggestions, and
to Mr. H. G. Newth for his assistance in preparing the paper for
press,
* Communicated by Prof. H. Maxwett Lerroy, M.A., F.Z.S.
47*
686 MR. P. R. AWATI ON THE MECHANISM
MaTEeRIAL AND METHOD.
I began my work in the Chelsea Physic Gardens of the
Imperial College of Science, but, since the specimens were scarce,
I had to go to Wisley, where I got good material through the
kindness of Mr. F. Chittenden, the Director of the Wisley
Horticultural Laboratory.
The following reagents were used for fixing the insects :—
(i.) Bouin’s solution,
(ii.) Carnoy solution (Formula No. IJ),
(iii.) Petrunkevitch solution ;
and all of these have given satisfactory results.
The insects were kept in the fixing reagent for 24 hours, and,
without being passed through the lower grades of alcohol, were
thrown into 90 per cent. alcohol, in which they were allowed to
remain for several weeks. Thus treated, they were sufficiently
hardened for section-cutting, and it was, therefore, not necessary
to keep them in absolute alcohol for more than three hours. After
immersion for. one hour in a mixture of equal parts of absolute
alcohol and chloroform, they were transferred to pure chloroform,
in which they were left for 24 to 36 hours. They were now
imbedded in paraftin-wax (melting-point 56° C.) in the following
way :—A saturated chloroform solution (in the cold) of wax of
the same melting-point was prepared and the specimens were
allowed to lie in it for two or three days at the temperature of
the laboratory. They were then transferred to pure molten wax
in the oven, where they remained for five hours. A block was
then prepared in the usual way.
The importance of attention to the details of the above routine
cannot be overrated. There is in the literature, so far as I can
find, no single detailed description of a method by which good
sections of hard insects can be got, and it is notoriously difficult
to find any but diagrammatic illustrations to the papers of those
who have investigated such forms. As the result, however, of
my experiments, | have found that it is possible to obtain series
of excellent sections, not only of Zygus, but of such hard and
thickly-chitinised insects as the bed-bug, without using any
reagent for softening the chitin. I am therefore justified in
giving some prominence to the technique.
Long soaking in a mixture of chloroform and wax was found
to be the only way to obtain proper impregnation, and the time
given is the optimum—a shorter time is insufficient, a longer
causes the tissues to become brittle. The length of the soaking
differs with different imsects, according to the quality of their
chitin.
The sections were stained, in the ordinary way, with Ehrlich
hematoxylin, orange G, and picric acid (saturated solution in
90 per cent. alcohol).
For macerating purposes, potash (10 per cent.) was used. A
few drops of acetic acid were found useful. The specimens were
OF SUCTION IN LYGUS PABULINUS. 687
generally cleared in turpineol, which did not make them brittle,
even if they were kept in it for a long time.
HoMo.LoGigEs oF THE MoutH-PArts oF RHYNCHOTA.
The problem of the true homologies is very difficult, and has
only recently been solved. In this section I propose to give a
summary of the literature dealing with this problem.
Fabricius gave the name “ Rhynchota” to the insects comprised
in the class “‘ Hemiptera,” on account of their sucking mouth-
parts. But till the time of Savigny (42), no attempt was made
to homologise their mouth-parts with those of other (biting)
insects. He hit upon the interpretation which is now generally
accepted, According to him, the mouth-parts of the Rhynchota
are homologous with those of the biting insects; the maxille,
the mandibles, and the labium being represented in these forms
by the internal stylets, the external stylets, and the proboscis,
respectively. His interpretation has been endorsed by Kirby
and Spence, Burmeister, and lately by Heymons and Leon.
Savigny and Cuvier, one with Wepa cinerea, and the other
with Ranatra linearis, had found the labial palps articulated to
the proboscis. But they could not discover any trace of the
maxillary palps, which are well-developed structures in the biting
insects. They had not, therefore, good evidence for the homologies
of the maxille.
These matters will be better elucidated if the history of the
homologies of each part be treated separately.
The Proboscis, or Schnabelscheide.
There were two views with regard to the structure of the
proboscis before 1880 :—
(i.) Burmeister (5), Latreille, and Graber held that the proboscis
was formed by the second maxille fusing together with the labial
palps. Burmeister has stated this view in very general and
vague terms, but it was Kraepelin (25, 26) who elaborated it in
detail. According to him the first segment of the proboscis
corresponds to the submentum and mentum together, and the
three segments, 2, 3, 4, to the three segments of the labial palps
of the biting insects. This view had been supported by the fact
that there were, on the second joint of the proboscis, certain
strong chitinous tubercles which were supposed by him to be
the rudiments of the organs, 7. e. extremities of external and
internal lobes of the palps. Leon (30), however, takes them
to be the -chitinous supports for the muscles of the proboscis,
running into the head.
This view has been exploded, and no present-day writers think
seriously of it, its interest being now purely historical.
(ii.) The second view was first formulated by Savigny and
Cuvier, and is now generally accepted. Savigny, in Vepa cinerea,
and Cuvier, in Ranatra linearis, discovered certain jointed
688 MR. P. R. AWATI ON THE MECHANISM
structures articulated to the proboscis, which they took to be
the labial palps. The palps are, therefore, in this view, separate
from the proboscis, and do not take any part in its formation,
the proboscis being formed by the fusion of the second maxille
only.
After Savigny comes Gerstfeldt (14). He elaborated this
view as Kraepelin had done the earlier one. According to him:
‘‘ Das erste Glied das nach Burmeister allein die Unterlippe
darstellt, wire an das submentum und entsprache den Cardines
der Lippenkiefer, das zweite Glied bestande aus den beiden
Stipites und wire analog dem Mentum, das dritte und vierte
Glieder gehérten zusammen den Endlappen der Unterlippe an und
entsprachen entweder nur den ausseren Laden (Paraglossx) oder
nur den unteren Laden (Ligulz), oder aber, was mir noch wahr-
scheinlicher ist, beiden mit einander vereinigten Ladenpaaren
zugleich.”
Leon (29-33) has fully dealt with the question of the homo-
logies of the proboscis. His examples are taken from the
Belostomide. He clearly demonstrated the presence of the labial
palps as distinct from the proboscis, and came to the following
conclusion, after examining all the examples :
“Hs ist ganz gleichgiiltig, in welcher Weise die Glieder der
Scheide von einer Art zur andern, sei es als Form, Grosse, als
Borstenanzahl, als Chitinerhebungen, etc. variieren mochten, eins
bleibt immer constant, das die Scheide aus derselben Zahl von
Gliedern besteht, die immer dieselbe Stellung zu einander haben
und die vollkommen homolog sind den Bildungsgliedern des
Labiums der beissenden Insecten.”
In the meantime, Dr. R. Heymons (18) had published the
result of his study of the development of certain rhynchotous
forms. His conclusions are:—(1) the labial palps have entirely
disappeared in the adult Rhynchota; (2) the so-called labial
palps are secondary structures from the third segment of the
proboscis ; they appear in the embryo but they degenerate and
disappear in the later stages. Leon, however, does not agree
with these conclusions. Heymons has deduced them from the
study of the comparative embryology of these forms; while Leon
has come to his different conclusions from the study of their —
comparative anatomy. His conclusions are :—(1) The labial palps
persist in the adult forms; (2) they have been discovered in
Nepa, Ranatra, and certain Belostomide ; (3) it is not possible
for any secondary structures to originate at the same place where
the primary structures had been before, and to perform the same
function as the latter.
While the homologies of the labium were being discussed in
Europe, a novel interpretation of the same structures was
put forward in America by Prof. J. B. Smith (45), who had
brought his special knowledge of the mouth-parts of Diptera to
bear upon this question. He held that the proboscis was a part
of the first maxille; the basal segment of the proboscis being the
cardo ; the second, subgalea; the third and the fourth, the two
OF SUCTION IN LYGUS PABULINUS. 689
segments of the galea. <All that remained of the labium was the
mentum—a boat-shaped process lying between the stylets. This
was an original view, confined to Smith alone. It was shown,
the next year, to be erroneous by Marlatt (34), and, after a few
years, by Meek (36), according to whom the boat-shaped process—
the mentum of Smith—was the pharynx.
Thus far no one has definitely demonstrated the presence or
absence of the labial palps. All the recent writers are unani-
mous in the view that the proboscis is formed by the fusion of
the second maxille, which consist of the submentum, mentum,
paraglosse, and ligule—the view held by Gerstfeldt long ago.
But here unanimity ends, and different views prevail as to the
existence of the labial palps.
(i.) Leon holds that they are present in the adult forms. He
deals with this question from the anatomical standpoint.
(ii.) Heymons holds that the labial palps appear only in the
embryonic stages, and disappear in the adult forms. He writes:
“Wenn einigen Autoren auch gewisse Anhange an der Unter-
lippe, also Palpi labiales angesprochen worden sind, so wird man
sich diese Deutungen gegeniiber skeptisch verhalten miissen, da
weder die Entwickelungsgeschichte, noch die vergleichende
Anatomie zu Gunsten solecher Ausnahmen sprechen.”
Maxille, or inner Borstenkiefer, or Stechborsten.
Savigny long ago, and others who have followed him, have
homologised the inner stylets of the Rhynchota with the first
maxille of the biting insects. But they have not clearly shown
what parts correspond to cardo, stipes, etc., and what becomes
of the maxillary palps—which are conspicuously absent in the
Hemiptera.
Kraepelin (25) held that the grooved inner stylets were
mandibles, which formed the tube for suction, while the maxille
were on the outside of the mandibles. This view is not now
accepted.
Huxley (48) denied the homologies of the inner stylets with
the first maxille of the other (biting) insects, because the former
do not happen to possess the maxillary palps of the latter.
Mecznikow (49) has also denied the homologies of the internal
and the external stylets in the Homoptera. According to
him the true maxille and mandibles appear in the embryonic
stages only, but they degenerate and disappear later on. Both
stylets of the adults are produced from the retort-shaped
(‘‘retortenformigen”) organs situated in the head. They are
therefore not homologous with the maxille and the mandibles of
the biting insects.
Witlaczil (50) has demonstrated, however, that the embryonic
mandibles and maxille do not disappear at all, but persist in
the adult stages, though there is a marked change in their
position. In early stages they are situated on the outside, but
later on they sink into the head and become entirely internal.
690 MR. P. R. AWATI ON THE MECHANISM
This sinking in, or involution, of these structures has been
brought about by the greater development (overgrowth) of the
clypeus and the labrum. The retort-shaped organs are nothing
but the swollen bases of these stylets. Not only are these organs
found in the Homoptera but they are also found in the Hetero-
ptera; though in the latter they are not so prominent as in
the former, owing to their degeneration. In the Homoptera
they have been recently demonstrated by Davidson (9).
The fact that both the stylets sink into the head owing to the
overgrowth of the clypeus, has been recently shown by Heymons.
The mandibles and the maxille, then, do not disappear but
persist in the later stages, though in different positions. ‘‘ Man-
diblen und Maxillenladen ziehen sich bei den Rhynchoten
in tiefe taschenformigen Hohlungen zuruck und scheiden die
chitinosen Stechborsten aus.” Thus writes Heymons. The
retort-shaped organs of Mecznikow, in the Homoptera, are the
‘‘taschenformigen Hohlungen” of Heymons, in the Hemiptera in
general.
Prof. J. B. Smith (45) was the first to call attention to the
fact that the maxillary stylets form but a part of the first maxille.
Each maxilla consists of two parts: the maxillary sclerite or
segment, and the maxillary stylet; but he was not able to
identify them separately.
It was not until Heymons had published his “ Beitrige zur
Morphologie und Entwickelungsgeschichte der Rhynchoten” (20),
that the relation between the maxillary segment or plate and
the maxillary stylet was clearly understood. According to Smith
the two pairs of stylets with the lateral (maxillary) sclerites
posterior to the mandibles, together with the proboscis, represent
the first maxille ; the stylets representing the lacinia and stipes ;
the sclerite representing the palpus and the proboscis. He had
made both the stylets arise from the same place. He is apparently
led astray in his interpretation by a faulty dissection, which was
shown to be the case by Marlatt (34).
Heymons is the first writer to explain clearly the homologies
of the first maxille. He has studied their development in certain
Hemipterous (both Heteropterousand Homopterous) forms. He
has clearly demonstrated that a maxilla arises as a single structure,
but that soon after it is divided into two parts :—(i.) a median
piece, or maxillenlade; (ii.) a lateral piece, maxillenhécker, or
maxillary plate (segment).
(i.) Lhe maaxillenlade.—This becomes elongated and transformed
into a long tapering stylet, two of which (one from either side)
combine together to form two tubes, one for suction of the sap,
and the other for ejection of the saliva.
(ii.) The maxillary plate—This represents the stem of the
maxilla (cardo and stipes) of the biting insects. It forms the
antero-lateral piece of the head-wall, and has therefore nothing
to do with the mouth-parts proper—except in so far as it forms
the support to the protractor muscles of the maxille.
OF SUCTION IN LYGUS PABULINUS. 691
The Maxillary Palps.
The presence of maxillary palps is one of the distinguishing
features of the first maxille of the biting insects, but they
have completely disappeared in the Rhynchota. The maxille
become elongated in these insects, and work inside plant or
animal tissues. Sensory apparatus in the form of palps is
superfluous.
Maxillary palps had been described long ago by Ratzeburg
(1827-34) in certain species. Hach palp was found to be three-
jointed. Burmeister (1835) had, however, found that they were
not palps but horny tubercles marking the attachment of the
maxillary muscles.
Heymons has, as far as possible, elucidated the question of the
maxillary palps. According to Smith, the maxillary sclerite and
the palps fused together to form one structure. Heymons,
however, holds another view. The maxillary palp is distinct
from the sclerite, and the maxillary plate has a process (processus
maxillaris) which he interprets to be the remains of a maxillary
palp. In some species of Tingide there are certain processes—
which were taken to represent the labial palps—but which are
now regarded as the maxillary palps.
Rudimentary palps are present in some Hydrocoridee—in
Nepa they are onion-shaped—but in Gymnocerata they are
identified with the Bucculz (?).
Mandibles, or external Stechborsten.
These were recognised as such long ago, though Kraepelin (25)
had mistaken them for the maxille, and had therefore made
them the sucking organs. The views of Mecznikow have been
given above. Prof. Smith, while working on the nymph of
Cicada, described a mandibular sclerite corresponding to that of
the maxilla, and a mandibular stylet also corresponding to the
maxillary. Heymons, too, has described the mandibular sclerite
(lamina mandibularis) and stylet in Cicada. The mandibular
sclerite is very well marked in the Homoptera, but in the
Heteroptera it is not present at all, or is rudimentary. These
structures are distinct in the later stages, though they have been
derived from the same structure in the embryo; the connection
between them is lost. The protractors of the mandibular stylets
are not attached to the mandibles but to their levers at one
end and, at the other, to the mandibular sclerite.
This interpretation of Heymons has been recently called in
question by Muir and Kershaw (22, 23, 24). The so-called
mandibular plates are not derived, according to them, from
the mandibles, and therefore have no relation whatever to
them. They are mandibular folds or sulci.
The most recent view therefore is :—(i.) The mandibles of the
biting insects are represented by the external stylets. (i1.) The
mandibles are not divided into two parts, one corresponding to the
692 MR. P. R. AWATI ON THE MECHANISM
cardo and stipes of the maxille and the other to the maxillary
stylet. (iii.) The protractor muscles of the mandibles are attached
to the levers and not to the mandibles.
Labrum.
This is the least disputed structure. Its homology with the
labrum of the mandibulate insects has been determined with
certainty. It is not, however, one of the appendages of the head
but a continuation of the clypeal sclerite, from which it is not
easily to be distinguished externally.
Tue Morro tocy or tHE Heap. (Text-fig. 1.)
The head of Lygus pabulinus consists of the following parts :—
(1) The Epicraniwm.—tThis lies between the two large eyes.
It is continued into
(2) The Clypeus. (The frons, the intermediate portion between
Text-figure 1.
Lygus pabulinus.
Mount of the whole head, macerated in potash (10 per cent.), and stained with
saturated picric in 90 per cent. alcohol; showing the internal chitinous
structures. Ob. 3 & Oc. 4.
The lines numbered 2, 4, 5 indicate the planes of the longitudinal sections
shown in the corresponding figures.
OF SUCTION IN LYGUS PABULINUS.
695
epicranium and clypeus, is not well marked.) The sides of the
clypeus have sunk into the head, forming clypeal folds to which
the protractors of the mandibles are attached, and which are
fused with the ventral wall of the pharynx (text-fig. 17, Clp.F.).
(8) The Labrum.—tThis is merely a continuation of the clypeus,
and there is no sign of external differentiation between the two
structures.
It extends as far as the first segment of the labium,
and tapers to a point, the base being broader.
It has rather a
deep groove underneath to hold the stylets in place, since the
labial groove at this point is too shallow and flat to do this.
There are three external longitudinal ridges on its dorsal surface ;
its sides are ornamented with small rounded lobes, and its surface
Explanation of Lettering.
. Antagonistic Muscles.
. Afferent Salivary Duct.
. Anterior Wall of the Pump-
chamber.
- Buccal Fold.
. Cambium.
. Circumesophageal Com-
missure.
p. Clypeo-labrum.
». Clypeo-labrum.
. Clypeal Fold.
. Clypeal Sclerite.
. Constrictor Muscles.
. Cortex.
. Cribriform Plate.
. Cardiac Valve.
. Dorsal Arms of the Ten-
torium.
. Divaricator Muscles.
. Ejection Canal.
. Epicranium.
Epidermis.
. Epipharynx.
. Endodermis.
. Efferent Salivary Duct.
», Gustatory Organs.
. Hypopharynx.
. Lumen of the A.S.D.
. Labium.
. Labial Groove.
. Labial Muscles.
r, Labrum.
. Ligament for A.M.
. Ligament for D.M.
. Ligament for P.M.
. Mandibles.
. Mandibular Articulation.
. Mandibular Lever.
. Protractors of the Man-
dibles.
. Retractors of the Man-
dibles.
. Cavity of the Mandibles.
. Hooks of the Mandibles.
. Mandibular Sheath.
. Mouth of the A.S.D.
|
Mx.C.
Mx.F.
Mx.L.
Mx.P.
Mx.R.
Mx.Md. Art.
Mx.Md.M.
Cavity of the Maxillary
Stylet.
Maxillary Fold.
Maxillary Lever.
Protractors of the Maxillary
Stylets.
Retractors of the Maxillary
Stylets.
Maxillo - mandibular Arti-
culation.
Maxsillo - mandibular
Muscles.
Sc. Maxillary Sclerite.
. Maxillary Sheaths.
. Maxillary Stylets.
. Operculum.
. (sophagus.
. Posterior Arms of the
Tentorium.
y. Pump-cylinder.
. Pump-chamber.
. Piston.
. Handle of the Piston.
. Pharyngeal Duct.
. Pharynx proper.
. Phloem.
. Pump Muscles.
. Retort-shaped Organs.
. Pump-stem.
. Posterior
Wall of the
Pump-chamber.
. Maxillary Process or Sup-
porting Strut.
. Suction Canal.
. Supra - esophageal Gang-
lion.
. Sub-cesophageal Ganglion.
. Salivary Duct.
. Body of the Tentorium.
. Trachea.
<. Tip of the Maxillary Stylet.
. Tip of the Mandible.
. “V” or ventral wall of the
Pharynx.
. Valvular muscles.
. Xylem.
694 MR. P. R. AWATI ON THE MECHANISM
is covered dorsally with small hairs or papillz, and is very smooth
ventrally. It is complementary in its function to the labium,
i. €. it is applied closely against the labium to form the deep
groove which keeps the stylets in place and prevents their lateral
movement (text-fig. 9, Lbr.).
(4) The Maxillary Sclerite—This is situated laterally to the
epicranium just below the eyes, and forms the lateral boundary
of the mouth. As stated above, it is one of the parts of the em-
bryonic maxilla. The protractors of the maxille areattached to it.
(5) The Labiwm (text-figs. 1, 29, Lb.).—This structure consists
of four segments. The first is broader and shorter and has
practically no groove, or, if there is any, it is very shallow and
flat. The first joint—i. e. the joint between the first and second
segments—is swollen, and this swelling is due to a great develop-
ment of chitinous ‘‘tendons” to which the labial muscles are
Text-figure 2,
7 (615 14:13
21 Ch. V. Mee wae Ay keSies
Nerve
to G Gre
Ph.
V. Ph.
Op.
Chitinous rods
in muscles
Many,
LUT
MUN
Mh
UD
ACT
Wo
“il
LUT
Gs. CV. Sub.G.
Lygus pabulinus.
Diagrammatic median longitudinal section. Ob. 3 & Oc. 4.
The lines numbered 9 to 22 indicate the levels of the sections shown in the
corresponding figures,
N.B.—In all the sections the microscope tube is not drawn out to its
proper length.
For explanation of the lettering see p. 693.
OF SUCTION IN LYGUS PABULINUS. 695
attached (text-figs. 2, 16, Lb.M.). Moreover, it acts as a hinge
upon which the whole of the labium is bent and doubled. This
is one of the characteristic features of the insect. In enables
the insect to get at the required tissues of a plant. The stylets
are thrust into tissues containing food, the depth of which below
the surface varies in different parts of the same plant. The
deeper such tissue lies, the greater is the bend of the labium at
the first joint. Its bend is scarcely appreciable when the insect
is sucking sap from tissues which are superficial.
The stylets cannot be increased in length, but this mechanism
by shortening the proboscis enables the insect artificially to
protrude them further into the plant tissue. The labial muscles
facilitate this bending (text-figs. 2, 16, Lb.M.).
Text-figure 3.
Pump-apparatus of Lygus pabulinus.
Sagittal section.
For explanation of the lettering see p. 693.
The second, the third, and the fourth segments are long and
narrow and gradually taper to a point. The groove, which is
shallow and flat in the first segment, begins to deepen in the
696 MR. P. R. AWATI ON THE MECHANISM
second, and, in the last segment, forms a tube enclosing the
stylets.. Cross-sections of the last segment show one tube within
the other. The inner tube enclosing the stylets is formed by the
groove; and the labium is the outer tube which encloses the
former (text-fig. 8, Lb.G.).
The. tip of the labium is encircled with bristles which are of
two sorts, fine and stiff. They are arranged in a definite way,
and function as sensory hairs. The insect feels the surface of a
leaf with them before thrusting its stylets into it.
The labrum and the labium together keep the stylets in place.
In the first segment, where the labial groove is too flat and
shallow to do it and where the labium hasa bend inwards, thereby
leaving the stylets free, the labrum encloses them in its groove.
Distally the labium takes over this function of holding the
stylets, since its groove becomes deeper and deeper, Thus the
stylets are always found in the groove of one or the other, and
are prevented from that lateral movement which would make
piercing and sucking impossible.
The labium protects the stylets in the groove, but its more
important function is very ingenious. The stylets are very thin,
needle-like structures which have, however, to pierce the tough
and cuticularised epidermis of a leaf. As they are very delicate,
they would bend in the act of piercing, were they not enclosed in
the proboscis. The lumen of the tubular tip of the proboscis is
so small that the stylets fit into it tightly (text-fig. 8). There
is no empty space in it for them to bend. The tip of the labium
is closely applied to the surface of the leaf; the protractor
muscles of the stylets contract; and the stylets are forced out
of the proboscis and driven against the epidermis, which they
cannot fail to pierce. Once in, their forward progress to the
required tissue is mechanical owing to the bend of the labium
at the first joint.
It is thus obvious that the labium is an important structure in
the sucking apparatus of this insect ; it is one, moreover, on the
structure of which stress has not been laid by previous writers.
(6) The Stylets, or Stechborsten. (Text-figs. 1, 6, 23.)
(1) MorpHonoey.
(¢) The Maxillary Stylets or Internal Stechborsten. (Text-
figs. 1, 5, 6, 23, Mx.St.)
It is shown above that the stylet is a part of the maxilla and
not the maxilla itself, Each maxilla is divided, in the embryonic
stage, into two parts, one of which forms the stylet or Stechborste.
The maxillary stylets are situated between the pharynx and the
mandibles on either side. They are not, however, on the same
level with either of them: they are just above the pharynx and
below the mandibles. They are hollow, and their cavities are
continuous with the body-cavity.
OF SUCTION IN LYGUS PABULINUS. - 697
Each of them consists ons
(i.) an external part, projecting beyond ite head and forming
the complementary half of the sucking tube ;
(ii.) an internal part, separate from its complemént and going
to one side of the pharynx.
(i.) The external part. This tapers to a fine and curved point.
In cross-sections it shows a groove which is divided into two
gutters by a longitudinal ridge which runs through it, from end
to end. In short, the whole structure looks in cross-section like
a W, and one groove is thus divided into two small, separate, and
non-intercommunicating groovelets. The two stylets from either
side come together and by approximation of these groovelets form
two complete tubes—one dorsal facing the labrum and functioning
Text-figure 4.
D.A.
Lygus pabulinus.
Nearly median longitudinal section, showing the supporting structure, the body
of the tentorium, the pump muscles, ete. Ob. 3 & Oc. 4.
For explanation of the lettering see p. 693.
698 MR. P. R. AWATI ON THE MECHANISM
as the suction-canal, and one ventral facing the labium, func-
tioning as the ejection-canal, through which saliva is forced down
by the salivary pump (text-figs. 8, 9, 10, Mx.8t., Su.C., E.C.).
There are not any extra-chitinous structures to unite the two
stylets to form the canals; the three ridges of each stylet
apparently uniting with those of the other. They can be detached
from one another with a little force. This fusion may easily be
seen in a series of cross-sections.
Text-figure 5.
Lygus pabulinus.
Lateral longitudinal section, showing the arrangement of the muscles
in the head. Ob. 3 & Oc. 4.
For explanation of the lettering see p. 693.
The tips of the stylets are smooth and lancet-shaped, as might
be expected from their function (text-fig. 23). In certain aquatic
forms they are ornamented with minute incurved hooks (0.
Geisse (18). Within the plant-tissues the tips of the maxille do
not end simultaneously ; one of them is pushed a little further
than the other (text-fig. 25).
(ii.) The internal part. This lies within the head. It gets
gradually flatter and broader when it is traced back. Hach stylet
OF SUCTION IN LYGUS PABULINUS. 699
ends internally in a swollen base. These swollen bases, which
are oval, have their importance in history, which has been referred
to above. They are known as the retort-shaped organs of
Mecznikow, or the Taschen-formigen organs of Heymons (text-
figs. 5, 6, 20, 21, Poc.Or.).
Text-figure 6.
Lygus pabulinus.
Reconstructed from numerous vertical sections, showing the relation of the
muscles to the stylets, the pump, etc. Ob. 3 & Oc. 4.
For explanation of the lettering see p. 693.
There is no trace of the maxillary groove upon the swollen
bases, but it begins a little below. The structure of the groove is
the same here as described in the external part, though on a
minute scale. The stylets are, of course, separate in the head, as
they pass one on either side of the pharynx (text-figs. 1, 5, 6,
11-19).
Proc. Zoot. Soc.—1914, No. XLVIII. 48
700 MR. P. R. AWATI ON THE MECHANISM
(6) The Mandibles. (Text-figs. 1, 5, 6, 23, Md.)
These are single structures from the beginning. ‘The embryonic
mandibles are the adult mandibles, there being no mandibular
sclerite. The present view of their homologies has been given
above.
Their external structure corresponds with that of the maxille.
They lie just above them and do not go so deep into the head.
Text-figure 7.
Lygus pabulinus.
Reconstructed from numerous vertical sections, showing the relation of the
muscles to the stylets, the pump, ete. Ob. 3 & Oc. 3.
For explanation of the lettering see p. 693.
OF SUCTION IN LYGUS PABULINUS. 701
They are flat and hollow and end in retort-shaped organs like
the maxillary stylets; but unlike them :—
(i.) They never form tubes, as they do not unite and have no
grooves on them. They are separate from end to end. Each of
Text-figure 8.
>_. _\ 5a
Lygus pabulinus.
Transverse section of the tip of the labium. Immersion-lens & Oc. 4.
For explanation of the lettering see p. 693.
Text-figure 9.
Lygus pabulinus.
Transverse section of the tip of the labium, showing the labrum and the labium
enclosing the stylets. Immersion & Oc. 4.
For explanation of the lettering see p. 693.
49*
MR. P. R. AWATI ON THE MECHANISM
Text-figure 10.
Lygus pabulinus.
A & B. Transverse sections, showing the epipharynx, gustatory organs, the
pharyngeal duct, the efferent salivary duct, etc. Ob. 6 & Oc. 4.
For explanation of the lettering see p. 693.
OF SUCTION IN LYGUS PABULINUS. 703
them is connected with the maxilla of its side by an interlocking
arrangement.
(ii.) The tips, which are pointed, are ornamented with recurved
hooks, the number and size of which seem to vary in different
species. The number varies from 7 to 9 in L. pabulinus. They
are bigger towards the tips of the mandibles but get reduced
posteriorly. The hooks will tear the tissues when withdrawn
from them with the maxillary stylets, though this arrangement
will not prevent their entrance into tissues. Their function will
be clear later on (text-fig. 23, T.Md.).
(II.) Anatomy.
The stylets are chitinous, the chitin being lined by the chitino-
genous epithelium which forms a continuous layer under it.
Kach of the stylets is enclosed in a sheath which is continuous
with the integument of the head. This shows that both stylets
have been invaginated in the course of their development, as is
demonstrated by Heymons (text-figs. 4-7, Mx.Sh., Md.).
The retort-shaped organs. (Text-figs. 5, 6, 20, 21, Poc.Or.).—
These are characteristic of the Rhynchota (Heteroptera and
Homoptera). Mecznikow (49) has described them as the structures
Text-figure 11.
, Lygus pabulinus.
Transverse section, showing thickening of the floor of the pharyngeal duct
(cf. figs. 10 A and 12).
For explanation of the lettering see p. 693.
from which the new maxille and the mandibles are formed, but,
as is shown above, they are not new structures at all. They are
nothing, in fact, but the swollen bases of the stylets, the swelling
being formed during development. They are the same as the
Taschen-formigen structures of Heymons.
They are oval and curved laterally outward. They are found
704 MR. P. R. AWATI ON THE MECHANISM
in the middle part of the head and show a curious structure in
cross-sections, appearing as areas of glandular tissue surrounded
by a chitinous ring which disappears in more distal sections.
The Interlocking Arrangement of the Stylets (external and
internal). (Vext-figs. 8-16, Mx.Md.Art., Md.Art.).—As shown
by Davidson (9), the maxille and the mandibles of Schizo-
neura lanigera are round and lie loose from each other. In
Lygus pabulinus, however, they are never separate outside the
head, but are interlocked. Within the head the members of both
pairs lie loose; the maxillee and the mandibles being smooth and
oblong without any processes (text-figs. 18-20, Md., Mx.St.).
Anteriorly in the labral region, they have a different structure.
The mandibles give rise dorsally to longitudinal ridges (which in
section appear as knobbed processes), which attach them to the
labrum or epipharynx. These disappear, however, beyond the
labrum. This arrangement demonstrates the importance of the
labrum in keeping the stylets in place and preventing their lateral
sliding.
The maxille also have small processes, of a similar nature and
function, which attach them firmly to the mandibles. These
structures are clearly visible in cross-sections. In the fourth
segment of the Jabium they are more pronounced.
It is obvious, therefore, that the mandibles and maxille are
inseparably attached to one another, and, the space between them
being exceedingly small, friction will ensure their simultaneously
working up and down; the mandibles accompanying the maxille
all the time and all the way in and out of the plant-tissues, unless
a differential force be applied *. Inside the head they are always
prevented from lateral movement by structures which are in-
vaginations or infoldings of the outer walls of the head, and
which enclose them successively at different levels.
These folds are formed by :—
(i.) The hypopharynx. (Text-figs. 15-15, Hyp.)
(ii.) The buccal fold. (Text-figs. 16-19, Bu.F.)
(iii.) The labial fold or the maxillary fold, formed by the
infolding of the maxillary sclerite. (Text-figs. 16-19,
Mx.F.)
These are very prominent structures in cross-sections of the
head. They are not, however, found in the heads of those forms
which require lateral movements of their mouth-parts, 7. e., in the
biting insects.
Tuer TENTORIUM OR ENDOSKELETON.
(Text-figs. 1, 4, 6, 7,17, 19-22.)
This has already been described by various writers: Burmeister,
Wedde, Léon, Bugnion, ete. Its description by Comstock and
* The existence of such a force will be demonstrated later on.
OF SUCTION IN LYGUS PABULINUS. 705
Text-figures 12 and 13.
Lygus pabulinus.
6 2AO ex: Se
ee SG
ee
Text-fig. 12.—Transverse section, showing the maxillo-mandibular articulation, the
mandibular articulation, the pump-chamber, the pump-stem, etc. Ob. 6 & Oc. 4.
6.St.
PSE.
Text-fig. 13.—Transverse section, showing the maxillo-mandibular articulation, the
mandibular articulation, the pump-chamber, the pump-stem, etc. Ob. 6 & Oc. 4.
For explanation of the lettering see p. 693.
Text-figures 14 and 15.
Lygus pabulinus.
Clp. Lbr.
Lig.
Md. Art.
Mx. Md. Art.
Mel Mr St
Ma.C. Md.
P St. Ee
Bu.F. S. SE.
PCA.
PCy.
FAG “y:
A.S.D,
Text-fig. 14.—Transverse section, showing the maxillo-mandibular articulation, the
mandibular articulation, the pump-chamber, the pump-stem, etc. Ob. 6 & Oc. 4.
Text-fig. 15.—Transverse section, showing the piston, the afferent salivary duct, the
divaricators, the protractors of the stylets, the labial muscle, ete. Ob. 6 & Oc. 4.
For explanation of the lettering see p. 693.
ON THE MECHANISM OF SUCTION IN LYGUS PABULINUS. 707
Kocchi (6) has the special advantage of a simple terminology,
which I have accordingly adopted here.
Text-figure 16.
Lygus pabulinus.
Transverse section, showing the piston, the afferent salivary duct, the divaricators,
the protractors of the stylets, the labial muscle, ete. Ob. 3 & Oc. 4.
For explanation of the lettering see p. 693.
The endoskeleton of Lygus pabulinaus consists of hollow chitinous
rods which are covered by a thin hypodermal (chitinogenous)
layer. They are nothing but involutions of the chitin of the
head, and therefore are called “ Apodemes.” The fact that the
tentorium consists of these hollow structures is very significant.
708 MR, P. R. AWATI ON THE MECHANISM
Great stresses have to be borne, and economy of skeletal material
has been effected and a maximum of rigidity attained, by forming
this material into hollow tubes instead of into solid rods. Many
parallel cases may be instanced from the plant kingdom—of
hollow stems which resist great stress.
Text-figure 17.
Lygus pabulinus.
Transverse section, showing the clypeal folds, salivary ducts. Ob. 2 & Oc. 12.
For explanation of the lettering see p. 693.
The tentorium consists of the following parts :—
(I.) The Body of the Tentorium, or Tentorium proper. (Text-
es. A. 7, 9) es)
This is formed by the fusion of the different arms of the endo-
skeleton. It lies between the pump below and the pharynx
above. It is found at the place where, if it were absent, there
would be great probability of the pharynx on the one hand and
the pump on the other, being dragged out of their positions
by contractions of their powerful muscles. It furnishes a firm
support for both these organs.
Its structure is interesting ; both its surfaces (dorsal toward the
pharynx, and ventral facing the pump) are curved and form two
grooves. The groove facing the pharynx is V-shaped and corre-
sponds exactly to the shape of the ventral wall of the pharynx,
which fits closely into it. They are joined together by con-
nective tissue.
The shape of the other groove, 7.e. the ventral one, is roughly
semicircular, and into it the pump is wedged, and kept there by
connective tissue.
OF SUCTION IN LYGUS PABULINUS. 709
Text-figure 18.
Ph,
Md Art.
-— Mx. F.
V. PA
Md. & Ma...
Mx. St. &
Mx.C:
Lygus pabulinus.
Part of a transverse section, showing the stylets magnified, and their
arrangement. Ob. 3 & Oc. 12.
For explanation of the lettering see p. 693.
(II.) The Dorsal Arms. (Text-figs. 4, 17, 18, D.A.)
These are short and do not reach the walls of the head but lie
free within it. The sides of the V (.e., the ventral wall of the
pharynx) are firmly attached to these arms by connective tissue.
(IIL.) The Posterior Arms. (ext-figs. 1, 4, 5, 7, 20-22, P.A.)
These are more important than the dorsal arms. They run
Text-figure 19.
Lygus pabulinus.
Transverse section, showing the mandibular levers, stylets magnified, and
their arrangement. Ob. 8 & Oc. 12.
For explanation of the lettering see p. 693.
710 MR. P, R. AWATI ON THE MECHANISM
posteriorly from the tentorial body to the posterior wall of the
head. They are broad, elongate, and flattened dorso-ventrally.
The pharynx is supported by them (text-fig. 20) just behind the
tentorial body, their connection with the pharynx being through
connective tissue.
At the far end, towards the wall of the head, the arms begin to
bifurcate, one armlet going straight to the posterior wall and the
other to the side of the head (text-fig. 5).
Their functions are very important in the mechanism of
suction :—
(i.) They support the pharynx at the place where there is great
tension.
(ii.) The pump muscles are attached to them ventrally.
(iu1.) The valvular muscles find support in them.
(iv.) The maxillary levers are supported by them.
Tt will be seen that the pharynx has support on all sides, 7. ¢.
ventrally, laterally, and dorsally (see below).
(IV.) The Levers. (Text-figs. 1,5, 6,19, 20, Mx.L., Md.L.)
Besides the tentorium proper, there are other chitinous struc-
tures which are connected to the outer and inner stylets. They
Text-figure 20.
Lygus pabulinus.
Transverse section, showing the antagonistic muscles, the retort-shaped organs,
the posterior arms of the tentorium, etc. Ob. 3 & Oc. 12.
For explanation of the lettering see p. 693.
are known as the mandibular and the maxillary levers, and are
placed at right angles to the direction of the stylets.
OF SUCTION IN LYGUS PABULINUS., ul
(A) The Mandibular Levers. (Text-fig. 19, Md.L.)
These are attached proximally to the lateral wall, or maxillary
sclerite, just below the bases of the antenne, and distally to the
mandibles through a igament. Their distal connection enables
the levers to pull down the mandibles when the protractor muscles,
Text-figure 21.
0.M._
Mx. St...
with
Mx.L.
Lygus pabulinus.
Transverse section, showing the maxillary levers, ete. Ob. 3 & Oc. 8.
For explanation of the lettering see p. 693.
which are attached to the former, contract. Proximally, at the
bases of the levers, there is no hinge-like device but, on the
contrary, they are fused with the sclerite. They are bifurcated
distally, the mandibles passing through the fork.
Their functions are :—
(i.) They give attachment to the protractor muscles of the man-
dibles, which muscles are attached to the distal ends only.
(u.) The maxillo-mandibular muscles are attached to them.
(Text-figs. 20, 21, Mx.Md.M.)
(ui.) The antagonistic muscles, which serve to bring back the
levers to their normal positions, are also attached to them.
(Text-figs. 5, 20, A.M.)
(iv.) The mandibles are supported by them.
(B) The Maxillary Levers. (Text-figs. 1,5, 21, Mx.L.)
These are narrow and elongate, and have no connection with
the lateral walls of the head. Their proximal attachment is to
the posterior arms of the tentorium, and distally they are con-
nected with the maxille. They taper gradually towards their
NY MR. P. R. AWATI ON THE MECHANISM
bases, and are curved as they pass below the maxillary stylets,
to which they are closely applied.
Text-figure 22.
Lygus pabulinus.
Transverse section, showing the supra-cesophageal ganglion, the retractor
muscles, etc. Ob. 3 & Oc. 8.
For explanation of the lettering see p. 693.
Their relations are :—
(i.) They support the maxillary stylets.
(1i.) The maxillo-mandibular muscles are attached to them.
(iii.) They never give attachment to any of the maxillary muscles,
which are attached to the maxillary stylets themselves.
Tue MUSCLES OF THE HEAD,
The musculature of the head is very complicated. The diversity
of the muscles is correlated with the diversity of function of the
structures which are required for effecting suction.
The groups of muscles are as follows :—
(1) The Divaricators. (Text-fig. 2, D.M.)
These muscles are attached to the upper wall of the pharynx ;
the mode of their attachment varying in different parts of the
head. In the clypeal region they are attached to the ligament of
the operculum (the upper wall of the pharynx); while in the epi-
cranium they are attached to the pharynx directly.
T find that these muscles are strengthened by chitinous rods
comparable perhaps to the tendons in the muscles of the verte-
brates (text-fig. 2).
OF SUCTION IN LYGUS PABULINUS. 713
These muscles fall into two groups, according to the mode of
their attachment and arrangement :—
(a) The fan- or feather-shaped muscles. (Text-figs. 14-21).—
These are arranged like barbson a feather vein or rachis, alternating
with one another on either side. This arrangement begins in the
clypeal region and gradually disappears posteriorly. They are
attached to the opercular ligament, which is the continuation of
the soft chitin lining the operculum. In the clypeo-labral region,
1.€.,1n the region where the gustatory organs are situated, the
ligament passes between those organs to give attachment to these
muscles. The gustatory organs consist of four lobes arranged in
pairs on either side of the dorsal wall of the pharynx. (Text-
figs. 27, 28.)
The fan-shaped arrangement is found in the region where there
isa great amount of work done; the work being to produce a
vacuum sutticiently powerful to suck in the sap which is being
accumulated in the pharyngeal duct. This vacuum is produced
by contractions of these muscles, the operculum being pulled ont
of the ventral wall of the pharynx. The arrangement described
gives a large surface for the attachment of the muscles.
(b) The strap-shaped muscles.—These are found in the epi-
cranial region, where they are directly attached to the upper wall
of the pharynx, which has here lost its character of operculum.
Here its ventral wall also loses its chitin, and the whole structure
is gradually transformed into the soft and thin cesophagus which
is no longer concerned with suction. These muscles are attached
to the dorsal wall of the head in the median line.
The divaricators occupy a large area in the head, beginning in
the clypeo-labral region and ending in the anterior part of the
epicranium.
(2) The Pump muscles, or Aspirators. (Text-figs. 2, 4, 5, 7,
17, etc. P.M.)
These are next to the divaricators in order of importance,
though they do not occupy such a large space. They are very
compact and powerful, as may be seen from the mode of their
arrangement and attachment. They lie below the pharynx in the
median line, but they diverge on either side towards the posterior
arms of the tentorium, to the ventral surface of which they are
fused. At the other end they are attached to the handle of the
piston of the pump.
When they contract they pull out the piston of the pump, the
posterior arms of the tentorium being very strong and rigid at the
sides. This pulling out cannot be an easy thing, small as the
pump is, because the posterior wall, of which the piston forms a
part, is made of thick, unyielding, though elastic, chitin, the
resistance of which must be enormous. Hence their dispropor-
tionate size and the mode of their attachment can easily be
understood.
14 MR. P. R. AWATI ON THE MECHANISM
(3) The Protractors of the Stylets. (Text-fig. 6, Mx.P., Md.P.)
(a) The Protractors of the Mandibles. (Text-figs. hes 19,
etc., Md.P.)
These muscles are attached at one end to the distal ends of the
levers of the mandibles (and not to the mandibles themselves), and
at the other to the clypeal folds, which run into the head to a
considerable distance. They are short and thick, and run obliquely
Text-figure 23.
|
|
Yd Se me Mx. St.
Md C
Md.H.
T. Mx.
T. Md.
Lygus pabulinus.
Tips of the stylets, showing the recurved hooks of the mandibles and smoothness
of the maxillary stylets. Immersion-lens & Oc. 4.
For explanation of the lettering see p. 693.
from one to the other. Their attachment at the distal ends of
the levers makes it easy to pull the latter down with the mandibles.
There is no hinge-like device at the bases of the levers, but when
they are pulled down they do not, I think, regain their normal
OF SUCTION IN LYGUS PABULINUS. 715
positions by their own elasticity alone or that of the side-walls of
the head. ‘They are brought to their original positions by the
antagonistic muscles.
The protractors are the chief agents in bringing about pro-
trusion of the mandibles.
(5) The Protractors of the Maxille. (Text-figs. 16, 19, 21, etc.,
Mx.P.)
They are attached at one end to the sides of the maxillary
Text-figure 24.
Lygus pabulinus.
Hand section, showing the styiets in the plant-tissue. Stained with saturated
picric in 90 per cent. alcohol. Ob. 3 & Oc. 4.
For explanation of the lettering see p. 693.
9 4S Now MUILIEX, 49
Proc. Zoou. Soc.
716 MR. P. R. AWATI ON THE MECHANISM
stylets (and not to the levers), and at the other to the maxillary
sclerite. It has been shown above that the latter is nothing but
the maxillary plate, the second half of the embryonic maxilla.
(+) The Retractors of the Stylets. (Text-figs. 16, 19, 21.)
These are attached to the sides of the mandibles and the
maxillary stylets just below the retort-shaped organs at one end,
and to the posterior walls of the head at the other. The stylets
show peg-shaped structures for the attachment of the muscles
Text-figure 25.
Md. —
= ae
Refer to pp.l2.
Lygus pabulinus.
Transverse section of a petiole, showing the maxillary stylets forward and the
mandibles behind, one of the hooks (*) of the latter being attached to the
cellular wall. Immersion lens & Oc. 4.
For explanation of the lettering see p. 693.
é OF SUCTION IN LYGUS PABULINUS. (Alii
(text-fig. 6), which are spread at their other ends over a large part
of the posterior wall of the head.
The retractors of the mandibles consist of two groups which
pass above and below the optic nerve (text-tig. 5).
(5) The Maxillo-mandibular Muscles. (Text-figs. 20, 21, Mx.
Md.M.)
These run from the mandibular levers to the levers of the
maxillary stylets on either side. It will be remembered that the
mandibles and the maxillary stylets do not lie at the same level,
one being above the other, and at the same time the maxillary
stylets project further back into the head than the mandibles. In
the normal position the tips of the mandibles and the maxillary
stylets are level, and in this position they are thrust into the
plant-tissue.
When these muscles contract, they pull in the mandibles and
push forward the maxillary stylets a little. Thus there is no
hindrance to suction by the maxille, as they are free from the
mandibles. At the same time the mandibles get themselves fixed
into the cellular walls by means of their recurved hooks and thus
steady the maxille for suction, as the latter are firmly attached
to the former by the interlocking device. Thus these muscles
play a useful part in the mechanism of suction.
(6) Zhe Antagonistic Muscles. (‘Text-figs. 5, 6, 20, A.M.)
These muscles are attached to the mandibular levers at one end
and to the postero-dorsal wall of the head at the other. They are
ealled antagonistic because they oppose the protractors of the
mandibles in their action, inasmuch as they bring the levers
back to their normal positions. They are very important, as there
is no other device to effect this readjustment.
They have nothing to do with the pharynx, as Bugnion and
Popoff (4) seem to think, who describe these muscles as ‘ Antagon-
istique ou Abaisseur du Pharynx.” The description that they
are attached to “une lame horizontale rattachée au Pharynx par
une expansion,” is absolutely inaccurate. This horizontal blade
(lame) is nothing but the mandibular lever, which has nothing
to do with the pharynx, but which is attached to the mandible.
‘Ta lame horizontale tendant a effectuer un mouvement de bascule,
Veffet de l’antagoniste doit étre d’abaisser le pharynx ou tout au
moins de le maintenir en place au moment oti le dilatateur entre
en action.” “La fonction du muscle antagoniste est de maintenir
Yappareil en place.” With due respect to these authors, I think
that their description is absolutely beside the mark. These
muscles have nothing to do with the pharynx. Wedde, however,
holds that there is no need for such antagonistic muscles of
Bugnion. He says, “‘ Antagoniste fiir die Schlundmuskulature
sind nicht vorhanden, es wirken als solche die héchst elastichen
chitinteile des Pharynx selbst.”
49*
718 MR. P. R. AWATI ON THE MECHANISM
Wedde, however, thinks that they are the clypeal muscles,
which have nothing to do with the mandibular lev ers, but whtteli
ave attached to them. He does not seem to have understood the
nature of the “horizontal bar,” which is nothing less than the
mandibular lever.
(7) The Constrictors or Circular Muscles. (Text-figs. 2, 7,
Con.M.)
These muscles are found in the cesophageal region. They lie
outside the external epithelium of the cesophagus. Their con-
traction and relaxation produce a kind of peristalsis which forces
the sap onward into the stomach. Bugnion (4) says, “ quant au
constricteur, son role doit étre de pousser dans lcesophage le
liquide absorbe.”
(8) The Valvular Muscles. (Text-fig. 7, V.M.)
They are short and thin and extend from the base of the cardiae
valve in the cesophagus to the posterior arms of the tentorium, to
both of which they ave attached. When they contract, the sides
of the valve are pulled apart, the valve thus opens, the lumen of
‘the csophagus widens, and the sap, which is under a great
pressure behind, is forced onward into the stomach, I cauinan
find them described by any previous writer.
(9) The Labial Muscles. (Text-figs. 2, 16, Lb.M.)
These are found in the first segment of the labium, and seem
to be important in increasing the amount of protrusion of the
stylets by bending the labium. They are attached at one end to
the supporting struts (buccal folds) on either side of the pump-
cylinder, and at the other to the first joint of the labium, which is
swollen, and thus offers a large area for the attachment of these
muscles.
It will be remembered that the proboscis bends upon itself, the
bending occurring at the first joint. The deeper the required
piant-tissue les, the greater 1s the bending of the proboscis.
There must be some device to effect this bending, and such
device is supplied by these labial muscles. When the muscles
contract, the first joint is pulled up and acts as a hinge—the
proboscis bending on it.
According to Geise, Wedde, and Nietsche they are elevators
and depressors of the labium. In the insect under consideration
no muscles are found that could possibly function in this way.
(10) Zhe Muscles of the Antenne.
As these have nothing to do with suction, they will not be
described.
N.B.—Wedde has described two pairs of protractor muscles of
the maxillary stylets, one of which is attached to the maxillary
OF SUCTION IN LYGUS PABULINUS., 719
sclerite, the other passing below the pharynx. It seems that
the latter pair does not represent the protractors but the pump
muscles.
Tur PHARYNX, on ScHLUNDKOPF. (Text-fig. 2.)
This is one of the interesting and characteristic organs of the
Rhynchota. In transverse sections it resembles roughly the
pharynx of a sucking Dipterous insect (Nuttall and Shipley (40)).
There are, no doubt, some differences in each case, but the general
plan is the same throughout. In all these cases suction is effected
by the production of a partial vacuum inside the pharynx, in
order to fill which the sap or blood flows up the proboscis.
The pharynx is a long, chitmous organ with a narrow
lumen. Its structure varies in different regions of the head, and
therefore it is proper to treat its several parts under different
names,
There are three distinct modifications in different regions :—
(i.) The Pharyngeal duct, in the clypeal region, where the
operculum as such does not exist, and the whole structure—here
formed from the lower wall of the pharynx alone—forms one
round duct which opens into the suction-canal. (Text-figs. 11-13,
PhD»)
(ii.) The Pharynx proper, in the clypeal region and the anterior
part of the epicranium, where its upper wall, as the operculum, is
clearly distinguishable. (Text-figs. 14-21, Ph.)
(ili.) The Csophagus, in the epicranial region, where the walls
of the pharynx are soft, and the operculum as such disappears.
(Text-figs. 2, 22, Cis.)
(i.) Zhe Pharyngeal duct.
This is the continuation of the pharynx proper into the labral
region where it opens into the suction-canal. It is the modified
pharynx, the modification consisting in a gradual elimination of
the epipharynx from the upper wall of the pharyngeal duct, from
behind forward. The lower wall at the same time gradually gets
thinner and more rounded and ultimately forms a duct by itself.
The divaricator muscles of the pharynx have disappeared with the
operculum. At the place where the pharynx ends and the duct
begins there are found the gustatory organs communicating with
the lumen of the duct through the ecribriform plate (text fig. 26,
G.Or., Cri.P].) In this region the hypopharynx (text-figs. 13, 14,
Hyp.) is well developed, forming a folded structure supporting
the maxillary stylets and the mandibles laterally, and enclosing
the efferent salivary duct underneath. It supports the pharyngeal
duct in the region where it has no other support, neither that
of the body of the tentorium nor that of the tentorial arms.
Anteriorly the hypopharynx becomes smaller and smaller and less
folded. It brings the efferent salivary and the pharyngeal ducts
nearer to each other (text-figs. 11-13, Ph.D., E.S.D.), and finally
720 MR. P. R. AWATI ON THE MECHANISM
disappears, leaving free these ducts, which open ultimately into
the suction- and the ejection-canals respectively (text-figs. 10 A
and 10 B).
The pharyngeal duct runs to a considerable distance into the
suction-canal before it ends, The importance of this arrangement
lies in the fact that it makes allowance for the movements—-up and
down—of the maxillary stylets. Had this duct run to a shorter
distance into the suction-canal, there would have been some
danger of its slipping out when the stylets were pushed down
to their natural limit of extension.
(ii.) The Pharynx proper.
This is found to begin in the clypeal region behind the clypeal
folds and to end in the anterior part of the epicranium, It is
bent anteriorly, and its bend corresponds to that of the head.
Posteriorly it is straight and runs directly into the csaphagus,
It is different in structure from the pharyngeal duct and from
the esophagus. It is the chief organ of suction in the insect.
When seen in transverse section it consists of two distinct parts :
the ventral part and the dorsal, The first may be called the “ V ”
and the second the operculum. ‘They are different in shape and
structure. (Text-figs. 14-21.)
(a) The “V” or ventral wall of the pharynx (V.Ph.).—This is
more or less V-shaped in section, its angle being considerably
drawn out in some places. In the anterior and the posterior
regions this angle gets rounder, and the characteristic form of the
“V" is lost, as in the pharyngeal duct and the esophagus. In
the clypeal region, where the pharynx proper begins, the arms of
the “ V” are elongated and fused with the clypeal folds running
into the head (text-figs. 16, 17, 19, Clp.F.). Thus it is dorsally
supported by these folds.
In the same region, moreover, the ‘‘ V” is wedged ventrally
into the body of the tentorium and retained there by the connec-
tive tissue, its ventral shape corresponding exactly to the dorsal
groove in the latter (text-figs. 17, 19, T.B., V.Ph.). The dorsal
arms of the tentorium run parallel with those of the ‘*V” and
support it laterally. It is this part of the pharynx which needs
support because it is here that the sucking force is applied,
The posterior arms of the tentorium as they run backward
also support the pharynx to some extent, but they soon diverge
laterally and leave it (text-fig. 20, P.A.).
The “ V ” is of thick chitin which, however, becomes thin and
soft anteriorly and posteriorly in the pharyngeal duct and the
cesophagus.
(b) Lhe Operculum or dorsal wall of the pharynx (Op.).—Its
structure is entirely different from that of the “V”. It is closely
apposed to the “V” in its normal position, which is regained
by its own elasticity, and from which it is pulled by contraction
of the divaricator (pharyngeal) muscles. It is obvious that there
OF SUCTION IN LYGUS PABULINUS. 021
is no space left in the lumen of the pharynx when the operculum
is in the normal position, but a strong vacuum is produced in it
when it is pulled out.
It is lined with a thin and soft chitin which is continued
dorsally into a ligament to which the divaricators are attached.
Posteriorly it loses its character as an operculum (text-fig. 22) ;
it is no longer introversible though it remains flexible. The “ V”
and the operculum form together, in this posterior part, one
structure, which is more or less circular and is continued in the
esophagus. In this region the muscles are directly attached to it
instead of to the ligament. As it passes anteriorly into the
pharyngeal duct it also loses its flexibility and introversibility.
It fuses with the epipharynx (labrum) which now covers the
ventral wall, and the muscles are again no longer attached to it.
Thus in both directions (anteriorly and posteriorly) the pharynx
proper loses its character as such, both in respect of the “ V”
and the operculum.
There is a hinge-like fold of chitin upon which the operculum
turns inside out. It will be understood from the figure (text-
fig. 20).
The supports of the pharnyx proper are :—
(i.) The hypopharynx.
(ii.) The clypeal folds.
(iii,) The body of the tentorium with its dorsal arms.
(iv.) The posterior arms of the tentorium,
All these supports make the “ V” immovably and firmly fixed
in its proper place, while the operculum remains all the time
flexible and introversible.
The function of the pharynx proper is very important since it
is the chief organ of suction. The pharyngeal duct cannot start
suction as there ave no muscles attached to its dorsal wall, which *
is no longer introversible; as explained above the operculum in its
normal position is closely apposed to the “ V,” and thus there is
no empty space between them. The muscles contract, the oper-
culum is pulled out, and a strong vacuum is produced. To fill it
up, the sap, which is now present in the pharyngeal duct, is sucked
into it,
Gil.) The Hsophagus.
There is no sharp external demarcation between the pharynx
proper and the cesophagus, the transition is gradual, the “ V ” and
the operculum losing their characteristic structures. In some
other species there is, at the entrance to the cesophagus, a constric-
tion, which is due to the fact that the thick chitin of the pharynx
proper ends suddenly where the cesophagus begins. ‘The nerve-
mass, consisting of the supra-csophageal ganglia, the lateral
commissures, and the sub-csophageal ganglia, surrounds the
esophagus, which then opens straight into the stomach through
the cardiac valve.
~I
BS
MR. P. R. AWATI ON THE MECHANISM
The csophagus is lined with the layers of an epithelium, and,
innermost of all,a soft chitin, which disappears posteriorly : from
this point, the outer layer of the cesophagus is generally one cell
thick, the cells being small. The inner layer consists of elongated
cells which have vacuoles in them, and which are greatly developed
in the stomach. Outside the outer epithelial layer are found the
vonstrictor and the valvular muscles. (Text-fig. 2, Con.M., V.M.)
The Cardiac Valve (text-figs. 2, 7, C.V.)—This is very short,
and it seems that its position in the cesophagus varies in different
insects. In Schizoneura lanigera it is present at the end of the
cesophagus (Davidson (9)), but in Lygus the valve is in its middle.
It is formed by the doubling of a part of the cesophagus upon
itself. The valvular muscles are attached to the base of the valve.
It remains closed in the normal position until the muscles contract,
when it is opened.
It prevents any return-flow of the sap from the stomach into
the cesophagus.
Tue Gustatory Oreans. (Text-figs. 26-28.)
That these are a specialised part of the supracesophageal
ganglia is proved beyond doubt. They are situated upon the
Text-figure 26.
Nucleus.
Nerve fibre.
Nerve from the brain Cribriform Plate
to the G.Or. with pores.
Lygus pabulinus.
Longitudinal section, showing the histological structures of the gustatory
organs with the cribriform plate, etc. Ob. 6 & Oc. 4.
For explanation of the lettering see p. 693.
operculum in the clypeo-labral region. There is a nerve running
from the brain to these organs on the upper wall of the pharynx.
Their histoiogical str uctures, though specialised, are similar to
those of the brain,
OF SUCTION IN LYGUS PABULINUS. G25
In vertical sections, they show four lobes, arranged in pairs on
either side of the operculum. The ligament passes through them
to give attachment to the divaricators (text-figs. 27, 28, G.Or.,
Lig.). There are nerve-fibres running into the pharynx, and
also from one lobe to the other. There are, moreover, small
nerves issuing from them and distributing themselves over the
epipharyngeal region.
The Cribriform Plate (text- figs. 27, 28, Cri.P].).— The
chitinous plate upon which these organs are situated is per-
forated, and through it the nerve-fibres communicate outward
Text-figure 27.
G.0r.
_ Nerve fibres joining
the lobes.
Upper wal! of —
the Pharynx.
Cribriform Flate-
Gils | tase
Lygus pabulinus.
Vertical section of the gustatory organs with the cribriform plate, showing
nerves passing to the periphery, etc. Ob. 6 & Oc. 4.
For explanation of the lettering see p. 693.
with the lumen of the pharynx. This cribriform plate seems to
be analogous with that found in the mammals, in which the
olfactory nerve-fibres pass through its pores to the olfactory
sense-organ.
These organs seem to be gustatory. Their function seems to
be to taste the sap as it comes into the pharyngeal duct.
724 MR. P. R. AWATI ON THE MECHANISM
Text-figure 28.
tig.
Upper wall of
the Pharynx.
Nucle/.
Nerve fibres
Joining the lobes.
Nerves to the
Periphery,
ee Soonr US.
Vertical section of the gustatory organs with the cribriform plate, showing
nerves passing to the periphery, ete. Ob. 6 & Oc. 4.
For explanation of the lettering see p. 693.
Tur PumMP-APPARATUS, OR WANZEN-SPRITZE.
This apparatus is one of the characteristic structures of the
Rhynchota, and has been described, though not in detail, by
many writers. It was’ first discovered by Landois (28) in
Cimex, and described later on by Wedde (46), in some: detail,
in Pyrrhocoris.
It has been found in almost all rhynchotous forms except the
Anoplura. Wedde has attached so much importance to it as to
divide the Hemiptera into two groups: (i.) Rhynchota setifera—
with the pump-apparatus, and (ii.) Pediculide—without it. It is
possible that, in the Pediculide, it may have escaped the notice
of investigators, since it must be very minute. I am the more
encouraged in this belief because Grove (16) says that the pump-
apparatus is not found in Siphonophora rosarum (a tolerably big
form), though he has figured it in his drawings (fig. 7 loc. cit.).
He further says, ‘‘In exactly the same position in the small
pointed under-lip.... which closes the mouth on its posterior
margin, where the above authors have described the salivary
pump, is a small U-shaped rod of solid chitin.” This small rod
of solid chitin is nothing but the pump-cylinder.
OF SUCTION IN LYGUS PABULINUS. 725
The pump may conveniently be divided into four parts for
description :—
(i.) The pump-cylinder with the pump-chamber and the
Bre ae) (Wexp-test 2.03 (7) [4015216 POys
-UD., £ OU.)
(ii.) The piston with the handle. (Text-figs. 15,16; Pi.,
lei dls))
(iii.) The efferent salivary duct. (Text-figs. 3, 10-15, E.S.D.)
(iv.) The afferent salivary duct. (Text-figs. 2, 3, 16, ete.
A.S.D.)
(i.) The pump-ceylinder with the pump-chamber and the pump-
stem.—The pump-chamber is more or less oval in section, and its
thick and rigid chitinous lining (constituting the pump-cylinder)
consists of two walls, an anterior and a posterior, which differ
from one another in structure and shape, though there is no
discontinuity of chitin between them.
(a) The anterior wall (A.W.P.). This is roughly semicircular
in section. Its middle portion is drawn out anteriorly into a
thickened solid process, which fuses with the hypopharynx, and
this thick portion of the pump is known as the pump-stem (text-
fig. 3, P.St.), which is perforated by the efferent salivary duct. The
chitin of the anterior wall is thick, elastic, and incompressible.
It gets, however, thinner laterally, and is continued into the
posterior wall dorsally; but ventrally it lines the entrance of
the afferent salivary duct.
(6) The posterior wall. This is apposed to the anterior wall in
the normal position. Its middle portion is swollen and is called
the piston. This is continued backward to form the handle.
Its chitinous lining is elastic, introversible, and can be retracted
from the anterior wall to a certain extent when the pump
muscles act. These muscles have already been described above.
(c) The pump-chamber. The space between the walls, anterior
and posterior, of the pump-cylinder constitutes the pump-chamber,
In the normal position, when the posterior wall is apposed to the
anterior, its capacity is reduced to a minimum. JIntroversion, of
course, increases this capacity. Into the chamber opens antero-
ventrally the afferent salivary duct, and from it issues antero-
dorsally the efferent salivary duct, which latter finds its way out
through the pump-stem to open into the ejection-canal of the
maxille. (Text-figs. 3, 10 B., P.Ch.)
(d) Attachment. The pump-cylinder is attached by connec-
tive tissue to the body of the tentorium, which is situated just
above it; the ventral groove of the tentorium corresponds
exactly to the dorsal semicircular contour of the pump-cylinder.
The attachment of the cylinder to the hypopharynx through the
pump-stem has already been mentioned under (a).
(ii.) The piston with the handle.—The middle portion of the
posterior wall is swollen and continued backward to a considerable
726 MR. P. R. AWATI ON THE MECIANISM
distance as a solid rod of chitin. To this the pump muscles
are attached. This structure constitutes the piston and the
handle, the swollen portion being the piston and the backward
continuation the handle. The latter is flattened dorso-ventrally.
(ii.) The efferent salivary duct.—This is a long and narrow
duct, issuing antero-dorsally from the pump-chamber. It has
a very narrow lumen and runs through the pump-stem and
perforates the hypopharynx before it opens anteriorly into the
ejection-canal. It has been shown above that the pharyngeal
duct also runs through the same region of the hypopharynx on
its way to the suction-canal. These ducts are thus brought
together and connected with one another by the hypopharynx.
Jt is the only connection between them; otherwise they are
distinct from each other. They do not communicate with one
another. And yet there are many misrepresentations about it.
Many of the previous writers are very doubtful of the facts, and
do not seem to know whether the efferent duct runs straight
into the ejection-canal or the labial groove, or opens into the
pharyngeal duct. Muir and Kershaw (22) write: ‘“ The syringe
or salivary pump. . . opens on the basal part of the labium
beneath the hypopharynx.” This statement is too vague to be
eriticised. According to them, it seems that the efferent salivary
duct opens into the labial groove. The same mistake has been
made by Grove (16), and is also found in many text-books. The
efferent salivary duct opens neither into the labial groove nor
into the pharyngeal duct, but, on the contrary, runs straight into
the ejection-canal and opens there. In short, the ducts, pharyn-
geal and efferent salivary, are separate from one another and
open into the suction- and ejection-canals of the maxillary stylets
respectively. (Text-fig. 10, Su.C., E.C.)
The efferent duct is supported by
(i.) The pump-stem, and
(ii.) The hypopharynx.
It runs a considerable distance into the ejection-canal before
it ends. This prevents its slipping out of the canal when the
maxille are pushed down by their protractors. (Cf. pharyngeal
duct.)
(iv.) The afferent salivary duct.—The two salivary ducts issuing
from the reservoirs on either side come together in the head and
form the afferent or common salivary duet under the brain.
This afferent duct is seen, in median longitudinal sections,
running underneath the pump-cylinder and. opening antero-
ventrally into the pump-chamber. It has a lumen bigger than
that of the efferent salivary duct, and its walls are flexible and
elastic. There is not a trace of a valve at its entrance into the
pump-chamber.
The salivary ducts (text-fig. 7, $.D.)—There are two salivary
OF SUCTION IN LYGUS PABULINUS, 127
ducts, one on each side, issuing from the salivary reservoirs in
the thoracic region. ‘They are narrow and Jong, and run straight
into the head, where they join together and form the common
salivary duct. They are always full of secretion, and as they
are thin and elastic, their walls are distended. The secretion is
therefore under pressure.
The Function of the Pump.
The structures of the pump, described above, will show that it
is used to force the salivary secretion forward into the efferent
salivary duct and thence into the ejection-canal of the maxillary
stylets. Hence it has been called a force-pump.
The ejection-canal pours the secretion straight into a wound in
a leaf made by the stylets. The secretion does not leak out since
the canal is air-tight, which will be seen from cross-sections.
The shape of the cylinder in different species depends upon the
degree of force required to drive the secretion through the
efferent duct. The more minute this duct the more cylindrical
is the pump: the return-stroke of the posterior wall is more
powerful in proportion to the force required to pull out the
piston. The posterior wall of the pump may be compared to
a bow.
In the normal position the posterior wall is apposed to the
anterior, and the capacity of the chamber is reduced to a
minimum. When the pump muscles, which are attached to
the handle, contract, the piston is pulled out and with it the
posterior wall. The capacity of the chamber slowly increases
and a partial vacuum is produced. Now, either the secretion
from the efferent duct or that from the afferent duct must flow
into the chamber to fill up this vacuum. The former case is
impossible, as the efferent duct is very short and empties itself
into the ejection-canal as soon as it is charged with the secretion,
and when empty its lumen would tend to collapse owing to the
thinness of its walls. On the other hand, the salivary ducts are
always full of secretion under pressure, and their walls are always
distended by their contents. As soon, then, as a vacuum is pro-
duced in the chamber, the secretion in the afferent duct flows
forward into it. Thus, every time the muscles contract and the
vacuum is produced, fresh secretion from the reservoirs flows into
the pump-chamber.
When the muscles relax, the posterior wall of the cylinder
begins to return to its former position, the return-stroke being
effected by its own elasticity. The fluid in the chamber 1s
gradually being compressed, and is forced along the line of least
resistance, which lies through the efferent salivary duct. This
duct is empty now, while the afferent duct is full of secretion
under pressure. The secretion therefore must flow into the
efferent duct. The space of the chamber is gradually being
reduced, and the entrance of the afferent duct into the chamber
728 MR. P. R. AWATI ON THE MECHANISM
is closed when that critical point is reached at which the pres-
sure inside the chamber becomes higher than that in the afferent
duct. This closure is effected by the posterior wall itself, which
slides over and closes the entrance to the duct. Thus there is no
necessity for a valve, and the saliva, once in the chamber, never
flows back into the afferent duct.
To sum up:—
(i.) The pump is a force-pump.
(ii.) The return-stroke of the posterior wall is effected by its
own elasticity, when it has been pulled from the anterior wall by
the pump muscles.
(iii.) The efferent salivary duct is short and empties itself into
the ejection-canal as soon as it is charged with secretion, It is
therefore empty when the return-stroke begins.
(iv.) The afferent salivary duct is always full of secretion,
which is always under pressure. It is longer than the efferent
duct.
(v.) No valve is required at the mouth of the afferent duct into
the chamber ; nor are there any muscles to open it. Its opening
and closing are automatic.
Tur MECHANISM OF SUCTION.
The functions of the different parts concerned in suction
have now been described under their respective headings. The
present section gives a connected account of the mechanism,
summarising, at the same time, what has been said above. The
following is a picture of the mechanism as it might be imagined
from the arrangement of the different structures described.
The mechanism consists of many structures in the dead co-
ordinated with one another, from the piercing of the epidermis
of a leaf by the stylets to the digestion of the sap in the stomach.
(1) The stylets inside a leaf.
The insect begins to feel the surface of a leaf with the sensory
hairs at the tip of the proboscis (text-fig. 29). It is trying to
find a good place for piercing. When it is found, the proboscis
is applied to it, the stylets are driven forward, the protractors
contract, and the epidermal layer is pierced, however thick it
may be.
(2) The bending of the proboscis.
Once in, the stylets are pushed down into the tissue by direct
mechanical force, 2.e. by the bending of the proboscis. This
feature has already been alluded to. It increases artificially the
protrusion of the stylets and enables them to reach the tissue
which eontains food. It seems that the insect is not confined to
any particular plant-tissues. The stylets may be found in the
xylem, the phloem, or in the cortex which contains food-substances
(text-figs. 24, 25).
“I
to
ble)
OF SUCTION IN LYGUS PABULINUS.
Text-figure 29.
The growwe of
the Proboscis.
3° Joint:
Sensory or
tactile hairs,
Tip of the
Frobosciss
Lygus pabulinus.
Mount, showing the labium macerated in potash and stained with saturated
picricin, 90 per cent. ale.
(3) Injection of Saliva into the wound.
When the stylets reach the particular tissues, the maxillo-
mandibular muscles begin to act. Their function has already
been dealt with. The mandibles, when they are pulled in by
these muscles, get fixed into the cellular walls by means of their
recurved hooks (text-fig. 25). The maxillary stylets are made
steady in them action by the mandibles because of their
interlocking arrangement.
Before the sap is sucked through the suction-canal, 1t seems
that saliva is injected into the wound made by the stylets. It is
possible that the saliva, by being mixed with the sap, may trans-
730 MR, P. R. AWATI ON THE MECIIANISM
form starch into sugar and thus help its digestion. Plateau (41)
has proved that the saliva of insects has this effect. It is
doubtful whether the sap is coagulated by exposure to air, but if
so, the saliva may prevent its coagulation. A third possibility
is that saliva may make the sap less viscid.
(4) Turgidity of the Cells.
This may be important indirectly in forcing the sap mixed
with saliva into the suction-canal of the maxillary stylets. The
cells of a tissue are always turgid and their walls are stretched to
their utmost capacity. They collapse if pricked, and squirt out
the sap with some force. The maxillary stylets pierce the cells,
the sap of which may thus be forced into the empty suction-
canal, since the other canal is full of the salivary secretion. I
attach no great weight to this possible factor in suction.
(5) Capillarity.
This factor may now come into action, and by its means the
sap would begin to ascend in the suction-canal if this were open
at the top. The sap reaches the pharyngeal duct, which hangs
into the suction-canal, and is thus immersed in the sap.
(6) Suction.
The divaricators of the pharynx begin to act. Those of the
anterior part of the clypeus contract first. The operculum is
pulled out and a vacuum is formed between it and the “ V.”
The sap is sucked into the pharynx from the pharyngeal duct.
The muscles relax, and the operculum regains its former position.
Meanwhile the muscles lying posteriorly contract, the operculum
is pulled out, and the sap is forced onwards. The muscles lying
still more posteriorly, do the same thing, and the same process is
repeated. Thus there is a wave of contraction of muscles passing
backwards, and the sap is continuously forced on towards the
stomach. It is prevented from flowing back, because there is no
empty space behind, since the operculum regains its normal
position with relaxation of the anterior muscles.
Thus, in the pharynx there are two complementary factors
which force the sap onwards towards the esophagus :—
(i.) The vacuum produced by the raising of the operculum.
(il.) Elasticity of the operculum, which enables it to regain
its normal position.
(7) The Gsophagus.
The sap is forced into the cesophagus, the walls of which are
soft and flexible and have attached to them the constrictor
muscles. As the sap distends the walls, the muscles contract,
and a peristalsis is produced which forces the sap onwards towards
the cardiac valve.
—
OF SUCTION IN LYGUS PABULINUS. 73
(8) The Cardiac Valve.
The valvular muscles contract, the valve opens, and the sap is
forced into the stomach.
(9) fhe Stomach.
This is a bag-like strueture and stores the sap.
To sum up :—
There are the following factors at work :—
G.) Capillarity in the suction-canal, helped by turgidity of the
cells.
(ii.) Suction produced by the vacuum through the raising of
the operculum.
(ii.) The peristalsis in the cesophagus by means of the
constrictors.
(iv.) The valvular action, which prevents the sap from flowing
back into the cesophagus from the stomach.
The pharyngeal duct hangs into the suction-canal and is
immersed in the sap. A question arises whether it is possible
for the sap to avoid the duct and flow into the body-cavity from
the suction-canal. The sap cannot do so because (i.) there is no
difference of pressure to force the sap into the body-cavity, and
because (ii.) there is active suction through the pharyngeal duct.
Another important thing for suction is that the suction- and
ejection-canals must be separate and distinct from each other.
The least intercommunication will stop the whole mechanism.
Also one canal cannot serve two contradictory purposes—one of
sucking and the other of ejecting saliva. The whole mechanism,
therefore, depends upon the presence of the two canals. (But,
curiously enough, the Bed-bug, Cimesx, is stated to present such
an anomalous case. It has only one canal for sucking the blood
and ejecting the saliva as well. Recently it has been described
by Dr. Max Braun (1): “In shape the maxille resemble two
gutters, the concave surfaces of which face one another and so
forma tube. This serves to conduct the blood from the wound
into the pharynx and also saliva into the wound. In other
Hemiptera food and saliva are conveyed by two distinct channels
formed by the longitudinal ridge which runs down the groove of
each maxilla, dividing it into two parts.” This is not the case
actually, as I hope shortly to show by figuring sections of its
mouth-parts.)
Damage to the Plant.
That Lygus pabulinus does some damage to potato plants
has been conclusively proved by the elaborate experiments of
Prof. Lefroy, of the Imperial College of Science, and Mr. Horne.
It is sufficient to say here that the insect drills into the leaves
holes which are bordered afterwards by brown rims. It does not
Proc. Zoou. Soc.—1914, No. L. 50
52 MR. P. R. AWATI ON THE MECHANISM
come within the scope of this paper to deal with the pathological
effects due to the insect-bites; but the way in which these holes
ave drilled is described below.
The structure and the shape of the stylets have been described
above. The tips of the maxillary stylets are smooth and lancet-
shaped, but those of the mandibles are deeply serrated (text-
fig. 23). The tip of the proboscis is covered with stiff hairs and
is blunt. How the stylets work inside the plant-tissue has been
described above. The insect is found on the same place for hours
at a stretch. It withdraws its stylets every few minutes from the
plant-tissues and thrusts them in again. ‘This process is repeated
many times. Since the mandibles can only be withdrawn by
tearing the cells each time they are taken out with their. recurved
hooks (text-fig. 25), and as this process is repeated many times,
a big area 1s “ultimately macerated and a hole is formed there
afterwards.
Tt was held by many writers—Dr. Riley being one of them—
that the Heteroptera made holes with the proboscis, the Homo-
ptera with the stylets. This view is absolutely inaccurate, since
it is found that the proboscis is incapable of drilling holes, its
tip being too thick and blunt to pierce the epidermis. Sections
of plants taken with the stylets iz situ show that the stylets
enter the tissues, the proboscis itself remaining outside, as
deseribed above.
BIBLIOGRAPHY.
(1) Braun (Max). 1910.—A Handbook of Practical Parasitology.
(2) Biseen (M.). 1891.—Der Honigtau. Jena. Zeitschr., Bd. xxv.
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(8) DANEDSON | (J.). 1913.—Biological Study of Schizoneura lanigera. Q.J.M.S,
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(9) Dimmockx. 1881.—The Anatomy of the Mouth-parts and the Sucking
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EO) ae oa: 1866.—Zur Anatomie der Hemipteren. Stettin. Entom. Zeit.,
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(11) Dreyrvus (L.). 1894.—Phylloxeriden. Zool. Anz. Bd. xvii. pp. 221-235.
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(18) Gets (O.). 1883.—Mundtheile der Rhy nchoten. Archiv f, Naturg.
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(14) ae 1853.—Ueber die Mundtheile der saugenden Insekten.
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(15) GrapeEr. 1877.—Die Insekten, Theil I. (Muiinchen.)
(16) Grovs (A. J.). 1909.—Anatomy of Stphonophora rosarum. Parasitology, .
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(17) Handbuch der Vergleichenden Physiologie, 1911. Bd. ii. Halfte i.. 2
(18) Heymons (R.). 1896.—Die Mundtheile der Rhynehota (Homoptera—
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a
OF SUCTION IN LYGUS PABULINUS. 733
Huymons (R.). 1897.—Ueber die Zusammensetzung des Insekten-Kopfes.
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Heymons (R.). 1899.—Beitrige zur Morphologie u. Entwickelung. der
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Kersuaw (J. C.) and Murr (F.). 1911.—Later Embryological Stages of the
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Leon (N.). 1897.—Beitrage zur Kenntniss des Labiums der Hydrocoren.
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Steiermark, Bd. xliv.
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Scumipr (E.). 1891.—Lippentaster bei Rhynchoten und Systematische
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- Weppke (H.). 1885.—Beitrage zur Kenntnis des Rhynchotenriissels. Archiv
f. Natureg., Jahre. li.
Wirraczit (E.). 1886.—Die Saugapparat der Phytophthives. Zool. Anz.
Bad. ix.
Houxtey (T. H.). 1858.—On the Organic Reproduction and Morphology of
Aphis. Trans. Linn. Soc.‘vol. xxii.
Mecznixow (N.). 1869.—Beitrage zur Embryonalentwickelunz der Insekten.
Troschel’s Arch. f. Naturg. Jahre. xxxv.
Wrrracatt (K.). 1882.—Zur Anatomie der Aphiden. Arb. a. d. Zool. Inst.
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Parasitology. vol. vi, No. 1.
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41. Procolophon trigoniceps, a Cotylosaurian Reptile from
South Afriea. By D. M. 8. Watson, M.Sc., F.Z.S.,
Lecturer in Vertebrate Paleontology in University
College, London.
[Received May 18, 1914: Read June 9, 1914. |
(Plates I.-ITi.* and Text-figures 1-5.)
INDEX.
Page
Structureweyei 7 ae ee eect | Yio
Systema ticpae yen cca eee erence) 6 HA:
The little reptile Procolophon has long been known, and the
work of Owen, Seeley, Woodward, and Broom has made us
acquainted with much of its structure. During my visit to South
Africa, I was fortunate enough to collect in the original locality,
Donnybrook, Upper Zwaart Kei, Dist. Queenstown, a nearly
complete skeleton (hereafter referred to as the female skeleton)
and several skulls and fragments. On the farm, Haslop Hill,
which lies about fifteen miles north of Donnybrook, I obtained
another almost absolutely complete and very well - preserved
skeleton, which I shall call the male skeleton.
This material, together with that in the British Museum, is
perhaps the most extensive in existence for the study of any
early reptile, and gives knowledge of almost every detail of the
structure.
Skull.—The general structure of the skull has long been known,
but it is now possible to deseribe the basis cranii and brain-case
in detail.
The basioccipital is a comparatively small bone whose posterior
end is formed by the transversely widened condyle, which is
rounded. The upper surface of the bone carries the exoccipitals
at the sides,-and the middle part of its upper surface is somewhat
depressed and supports the brain. On the ventral surface the
bone is soon covered by the basisphenoid but contributes to
the tubera.
The basisphenoid is a large and remarkable bone; on its lower
surface two strong ridges continue the tubera forward and termi-
nate in the basipterygoid processes, whose flat articular faces look
well to the front. It is possible that the basipterygoid process
is pierced by a foramen Vidii, but this is not certain. Between
these ridges is a very deep groove, which in front passes upwards
into the skull, separating the basipterygoids. From the sides of
the bone immediately below and in front of the fenestra ovalis, a
short but relatively powerful process passes outwards and back-
wards and ends freely. On the upper surface the basisphenoid is
a
* For explanation of the Plates see p. 747.
736 MR. D. M. S. WATSON ON
much excavated in the middle, and its lateral borders rise to the
opisthotics. The upper surfaces of the processes spoken of above
are deeply grooved, and the grooves from them pass on to the body
of the bone, forming a depression stretching right across it. In
front of this is a ridge which marks the posterior border of the
pituitary fossa, whose surface is not very well preserved in any
case.
Text-figure 1.
BSP Vest. BOc.
Procolophon trigoniceps.
A. Base of skull and otic region viewed from below and the right side. X 2.
B. View of lett side of brain-cavity. 2.
B.Oc., Basioccipital ; B.Pr., Basipterygoid processes ; B.Sp., Basisphenoid ;
Ev.Oc., Exoccipital ; F.Jug., Foramen jugulare; #.V., Foramen Vidii? ;
Fen.Ov., Fenestra ovalis; Op.O., Opisthotic; P7.0., Pro-otic; Vesti.,
Vestibule.
The exoccipital is a bone of medium size, articulating with the
upper surface of the basioccipital. It is comparatively long, and
is pierced at about the middle of its length by two small foramina
for the twelfth nerve, which he just above its lower border.
These two openings soon join and open ina single foramen on
the outside of the cranium. The bone extends outwards for a
little distance with its upper part in contact with the posterior
surface of the paroccipital, from which bone it is separated below
by the large, round foramen jugulare. The exoccipital forms the
A FOSSIL REPTILE. (avi
side-wall of the foramen magnum, and its upper end is in contact
with the supraoccipital.
The opisthotic is a large and powerful bone. Its inner end
was largely cartilaginous, and its ossified part does not touch the
basis craniil. Part of its posterior face is covered by the ex-
occipital, lateral to which it forms the very deep and massive
paroccipital process, the distal upper corner of which is connected
with the tabulare. The front of the paroccipital process is
covered by the pro-otic, and the two bones combine to form a
rounded groove on the lower surface in which the stapes lies.
The pro-otic is applied to the anterior face of the opisthotic,
and resembles that bone in having a largely cartilaginous inner
end. This is, however, larger in area and is clearly pierced by a
foramen, or possibly two, for the VIIthand VIIIth nerves. On
the inner side of the skull the inner ear is widely open and lies
high in the side-wall of the brain-cavity. The whole condition is
extremely like that in a tortoise. The front edge of the pro-otic
has a large incisura for the trigeminus, below which is a pro-
minent process. The fenestra ovalis is a very large irregular
hole in the bony skull.
Text-figure 2.
ja
EZ Zi) =
Procolophon trigoniceps.
A. Posterior aspect of skull. 1.
B. Right lateral aspect of skull. X 1.
Composite figures from three perfect skulls.
The supraoccipital is a very delicate bone, whose inner aspect is
not known; externally it is exactly like that of a lizard, touching
the parietals by a very small point, if at all directly.
The stapes is a relatively thick rod, whose outer end is expanded
into a plate.
There is a slender epipterygoid, which expands below and is
broadly fastened on to the outer surface of the pterygoid just
738 MR. D. M. 8. WATSON ON
outside its articulation with the basisphenoid. This rises in the
skull till its upper end nearly touches the parietal lateral to the
parietal foramen.
The general structure of the roof of the skull is already quite
familiar from the figures and descriptions of Seeley, Smith
Woodward, and Broom. The figures in this paper will, I hope,
explain any doubtful points: the relations in important regions
are described below.
The quadrate is a comparatively small bone; it has a small
condyle for the mandible, above which it rises vertically. The
body of the bone is in contact with the very large and massive
quadratojugal at its lateral border; it is then separated from
this bone by the quadrate foramen. Above this level the whole
posterior surface of the quadrate is covered by the squamosal.
The quadrate sends a long process inwards which is covered
posteriorly by the posterior ramus of the pterygoid.
The squamosal isa bone of medium size which may be described
as consisting of three parts. The posterior and largest of these
covers much of the back of the quadrate; it curves round to
form the second region, which lies entirely on the outer side of
the skull, having sutures with the quadratojugal, jugal, post-
orbital, and tabulare. The third part arises from the other two
and passes inwards and backwards over the otic notch to nearly
or quite touch the parietal. The whole of this part is covered by
the tabulare.
The tabulare is a very large bone forming the posterior corner
of the flat upper surface of the skull. Its anterior end overlaps
the parietal and squamosal to a very large extent, but its hinder
margin is free, and is deflected to meet the upper posterior corner
of the paroccipital.
Of postparietals there is usually no trace, but in the skull of
the female skeleton, which is well preserved, there are quite
clearly a pair of small bones articulating with the pointed mesial
posterior ends of the parietals. These are very small, only about
1 millimetre square, but can only be interpreted as vestigial post-
parietals. They stand out quite freely and touch no underlying
bones.
An interesting point is that the nasal cavity is very largely
separated from the very large orbit by plates arising’ from the
lachrymal and prefrontal.
The septomaxilla is a small bone articulating with the pre-
maxilla and nasal and forming the posterior border of the nostril.
‘The material at my disposal has shown no trace of a para-
sphenoid, but Dr. Broom has recorded it in an Albany Museum
specimen.
A curious feature of the very well-preserved skull of the male
skeleton is the presence of a small depression with rather stee
walls on the outer surface of the maxillaand nasal, just behind the
upper end of the septomaxilla. This prabably lodged a superior
labial gland.
A FOSSIL REPTILE. 739
One feature of the skull about which there has been some
dispute is the presence or absence of a foramen in the temporal
region. There is no doubt that Broom is quite correct in denying
its, presence, but in justice to Prof. Seeley and the British Museum
mason, it should be pointed out that the hole in this region in
their skulls i is not an artifact, but is due to a rotation inwards of
the posterior edge of the postorbital as a result of slight
crushing.
Text-figure 3.
Procolophon trigoniceps.
(A) Outer, and (B) Inner aspects of lower jaw. X 1.
Ang, Angular; Cr., Coronoid; Den., Dentary; P.Art., Prearticular ;
Sp., Splenial; Sur. Ang., Surangular.
Composite figures from two specimens.
The structure of the lower jaw will be understood from text-
fig. 3. The sutures separating the prearticular from the articular
and angular are not clear in any specimen, but I think they are
faintly shown as figured in the skull of the female skeleton.
Every other suture is obvious.
The coronoid extends forward between the dentary and splenial
for some distance.
Vertebral column.—There are twenty-six presacral vertebre.
The axis has a powerful spine, very long anteroposteriorly ; and
powerful posterior zygapophyses, which are continued forwards
into a broad table. There is a large intercentrum between the
centrum of the axis and the odontoid. The odontoid is not known
in detail.
The atlantal intercentrum is large and its anterior face forms
part of the cup for the occipital condyle. There is a pair of
neural arches to the atlas. These are separated posteriorly by
the neural spine of the axis; each has a facet on its inner face by
which it articulates with the odontoid, anteriorly it has an
articular face for the basioccipital. In front its upper surface is
overlapped by the proatlas, and behind it articulates with the
740 MR. D. M. S. WATSON ON
prezygapophysis of the axis. The atlas is very short from back
to front. The proatlas is paired (and articulates with the skull ?).
The remaining presacral vertebre are all much alike, but their
neural arches become heavier as they are traced backward. The
centra are completely pierced by the notochord and are consider-
ably constricted, the lower surfaces, however, being flattened.
The articular ends are expanded. There are intercentra between
all the presacrals ; these are single crescentic bones of an ordinary
character. Broom records paired intercentra between the third
and fourth vertebree. I have not seen this region.
The neural arches are heavy, extremely so in the posterior
part of the column. The articulating faces of the zygapophyses
are flat and placed horizontally throughout. The neural spines
are only represented by very short projections on the top of the
swollen neural arch.
The rib articulations are single, carried at the end of a short
transverse process, contributed to by both arch and centrum, in
the anterior part of the column; with the progressive increase 1n
size of the neural arch this process becomes obsolete, but the rib
articulation remains essentially similar.
The first sacral vertebra is very short anteroposteriorly ; it
carries a very large rib articulating with both arch and centrum,
and its neural arch is heavy. The anterior zygapophyses are
wide apart, but the posterior facets are placed close together.
The second and third sacral vertebre have slender arches, but
their articulation-faces are still flat and horizontal. The caudal
vertebre are not well exposed in any specimen ; anteriorly they
resemble the posterior sacrals, but further back they develop
fairly high spines and long chevron bones. It is not known how
far back they carry ribs.
The ribs are single-headed throughout and present no feature
of interest. There is some evidence of a rib on the atlas, and
there are certainly ribs on all the other presacral vertebra except,
perhaps, the last two or three, on which I have not seen any;
even in a well-preserved skeleton, if present, they were short.
Abdominal ribs.—Three specimens show the whole series of
abdominal ribs. The plastron is extremely reduced, consisting of
at most five series of transverse rows each composed of six very
small slender elements. These are divided into two groups, which
do not meet in the middle line. The whole arrangement is about
a centimetre from back to front, and lies about 15 mm. behind
the pectoral girdle, cf. text-fig. 4, B.
Pectoral girdle—The general features of the pectoral girdle
have long been known.
The scapula stands nearly vertically and is a simple bone with-
out an acromion. It has a ridge down its outer surface and
thickens considerably at the glenoid cavity. The anterior part of
the lower margin of the bone is in contact with the precoracoid,
and for the anterior half of this union the external surface of
one bone is directly continued on to the other. Posteriorly this
A FOSSIL REPTILE. 741
surface is interrupted by the notch which forms the glenoid
cavity. The posterior part of this is formed by the coracoid,
which articulates with the scapula and precoracoid, and extends
backwards behind them for a long distance.
The glenoid cavity thus formed by all three bones looks almost
directly outwards and is essentially a trihedral pit; it must have
been covered during life by a thick pad of cartilage, but can never
have shown any trace of the screw-shape shown in early Coty-
losaurs such as Diadectes, Labidosaurus, and, to a less extent,
“ Pariasaurus.” .
Text-figure 4.
Procolophon trigoniceps, 6.
A. Right lateral aspect of shoulder-girdle. X 1.
A’. Ventral aspect of shoulder-girdle. X 1.
B. Ventral aspect of shoulder-girdle, and abdominal ribs. X 1.
C. Right side of pelvis from outside. 1.
The so-called notch in the precoracoid proves when fully
exposed to be really a foramen passing through the bone and
opening on its upper surface. Its most remarkable feature is
the strong rounded ridge which forms its anterior border on the
lower surface.
The clavicles are large bones, beginning at the upper margin
of the scapula; they run down the whole anterior border of that
bone and then turn suddenly in, being vertically expanded and
applied to the anterior face of the T-shaped interclavicle.
The humerus of Procolophon is a very remarkable bone. The
742 MR. D. M. S. WATSON ON
head is a large triangular area, obviously covered by a thick pad
of cartilage during life.
The anterior lower corner of the head gives rise toa ridge
which rapidly subsides into the shaft and forms the only repre-
sentative of the deltoid crest.
The anterior upper corner also gives rise to a ridge which
passes down the dorsal and anterior edge of the humerus until it
in turn passes into the shaft.
The posterior lower corner of the triangular head gives rise to
a pronounced ulnar crest of much greater size than the radial
expansion. The bone has a narrow shaft which expands into the
wide lower end of the bone. This bears a prominent condyle for
the head of the radius and one for the ulna. There is a large
entepicondyle with a foramen.
The radius and ulna present no marked features.
The carpus and manus are exactly as Dr. Broom has already
figured them; the radiale is unossified; in the male skeleton
traces of its cartilage were preserved.
Pelvic limb.—The pelvis, as shown in my specimens, is rather
different from that of Dr. Broom’s specimen.
The ilium is high and narrow, it is supported by the three
sacral ribs, and placed almost vertically; it forms about a third
of the acetabulum, above which it is thickened to form the usual
process.
The pubis is an extremely massive bone, nearly square when
seen from below, and pierced by the foramen ; it articulates with
its fellow so that the lower surfaces meet in an angle considerably
less than two right angles; in other words, the lower surface of
the pelvis is not flat. The upper surface of the pubis slopes
down very rapidly in front.
The ischia present no special features.
The hind leg has already been well described and figured by
Seeley and Broom; my specimens agree exactly with theirs.
The restoration of the skeleton in a walking attitude, text-
fig. 5, is drawn entirely from the male skeleton.
The length of the neck depends on the actual position of the
pectoral girdle in the specimen, which is quite undisturbed, lying
on the ventral surface with the fore legs placed straight back
along the body and the soles of the two feet together along the
tail. The head is turned sharply to the right, but as every bone is
in close and accurate articulation this position must be a possible
one. In the female skeleton the head is in a similar position,
and the shoulder-girdle equally far back. The tail in the
restoration is completed from the entire series of caudals of the
female skeleton.
Taken as a whole Frocolophon is very lizard-like in build,
although it is rather massively constructed.
The male and female skeletons are of exactly the same length,
as are their skulls. The dentition is quite similar in the two
specimens, and I think there is not the slightest doubt that they
A FOSSIL REPTILE. 743
belong to the same species. The female skeleton is, however,
much more lightly built, having narrower and more slender limb-
girdles, and limbs which are less than three-quarters as long as
those of the male skeleton and much less robust. Mr. Boulenger
tells me that differences of this kind often distinguish the
Text-figure 5.
Procolophon trigoniceps.
Restoration of the skeleton of a male, all the bones except those in dotted
outline being drawn from a single individual. X 2.
744 MR. D. M. S. WATSON ON
sexes in living Lizards, and I think it is extremely probable that
the same reason accounts for the difference between my two
skeletons.
Affinities of Procolophon.
That Procolophon is a Cotylosaurian has been recognised by all
recent workers. Its right to inclusion in this group depends
on its roofed skull (it is perfectly obvious from the examination
of well-preserved skulls that the large opening on either side is
entirely an enlarged orbit and not, as has sometimes been sug-
gested, a joint orbit and temporal fossa), and on the typically
Cotylosaurian vertebre which are very similar to those of
Labidosaurus.
Mere reference of an animal to the Cotylosauria means very
little, as that great group includes animals which differ at. least
as much from one another as do a Lizard and Sphenodon, or even
a Tortoise and a Crocodile.
When discussing Pariasaurus recently I divided all Cotylo-
sauria, after the removal of the very primitive Seymouria, into
two groups, from the way in which they obtained a vertically
standing quadrate, one process leading to the obliteration of the
otic notch, the other to its exaggeration.
To the first group belong the Captorhinids, Pariotichids, and
Pantylids; to the second, Diadectids and Pariasaurids.
Procolophon has an enormous otic notch, and of course falls
into the second group.
Procolophon and its allies are the latest of all Cotylosauria,
occurring, so far as is definitely known, only in the Trias after
the disappearance of all other types. In consideration of their
age we should expect to find many advances in their structure.
[Any study of paleontology shows that all allied animals tend,
with mere passage of time, to change their structure in certain
definite ways which are common to whole groups and occur
independently in different lines of descent. These changes,
which are usually independent of the adaptations of the animals
in which they occur, take place at different rates in the distinct
lines, and the sets of changes of different parts of the same
animal are independent and may proceed at very different rates.
Changes of this type I refer to as advances; they are quite
different from specialisations, which themselves are of two kinds,
adaptations to some special mode of life, and what are usually
regarded as specific characters, 7.e. such differences between
animals as form the typical Mendelian allelomorphs; it is very
probable that these characters once established, perhaps as salta-
tions, are very stable, and may persist in a recognisable form
during a long series of advances. |
The advances in the structure of Procolophon are :—
1. The rounded occipital condyle.
2. The vertical quadrate.
A FOSSIL REPTILE. Ae
3. The loss of the primitive connection between the squa-
mosal and pterygoid.
The loss of the ‘‘ supra-” and inter-temporal bones.
The reduction of the lachrymal so that it no longer
reaches the nostril. i
The release of the distal end of the stapes from the
quadrate.
7. The short atlas and long axis.
8. The long neck.
9
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9. The extreme reduction of the plastron.
10. The loss of the cleithrum.
11. The loss of the supraglenoid and glenoid foramina.
12. The loss of the screw-shaped glenoid cavity.
13. The loss of the fifth distal carpal.
14. The loss of the centralia tarsi and the fifth distal tarsal.
The specialisations in the structure of Procolophon are :—
1. The enlarged tabulare.
The enlarged quadratojugal.
The enlarged orbit.
The dentition.
The characteristic humerus.
The non-ossification of the radiale.
The three sacral vertebre.
MI OU HE Oe bo
No other Cotylosaurians approach Procolophon in the number
of its advances, no other form has the lachrymal excluded from
the nostril, and none other has completely lost the screw-shaped
glenoid cavity. These differences, however, are not of such a
character as to preclude the possibility of Procolophon having
been derived from some known earlier form; they are merely
a result of its lateness in time.
The very large otic notch seems by itself to render any con-
nection with the Captorhinide impossible; but of even greater
importance is the difference in the brain-cases. Procolophon has
the vestibule of the inner ear occupying the whole side-wall of
the cranium, and in wide connection with the brain-cavity, exactly
as ina Tortoise or Sphenodon. The Captorhinids, although the
material is not sufficiently good for certainty, seem to have the
vestibule placed very low down, as in a Therapsid; a difference
of this character seems to me to render any idea of a close
connection between the groups quite unbelievable.
So little is known of Pariotichus and Pantylus that their
affinities are quite uncertain. |
Pariasaurus differs from Procolophon in its very long, low
brain-cavity, with a bony separation of the inner ear from the
cranial cavity. It is, of course, a very much less advanced type,
retaining a concave occipital condyle, a trace of the connection
of the squamosal and pterygoid, a lachrymal reaching the nostril,
TAG. MR. D. M. S. WATSON ON
a cleithrum, a screw-shaped glenoid cavity, and a fifth carpal.
‘A blood relationship between Procolophon and Pariasaurus is
more probable than one between Procolophon and Labidosaurus,
despite the very different specialisation of the two types.
Diadectes resembles Procolophon in its large otic notch, and
apparently, to some extent, in the features of the cranial cavity.
It differs, however, in the complete closure of the post-temporal
fossee by a union of the widened supraoccipital with the tabulares
and postparietals. In addition to these phylogenetically im-
portant differences are many due to the very much less advanced
nature of the type.
Limnoscelis differs in the apparent obliteration of the otic
notch and in its closed post-temporal fossz, in addition to the
retention of many primitive structures.
The work of Boulenger and v. Huene has shown the very
great resemblance between Telerpeton and Procolophon, and I
have been able to confirm their accounts on a magnificent new
skeleton of the latter lent to me by Mr. Taylor of Llanbryde.
Dr. Broom, when he first examined Procolophon, expressed
his opinion that whilst technically a Cotylosaur it was really a
primitive “ Diaptosaurian,” picking Paleohatteria as perhaps its
nearest known relative. This choice was rather an unfortunate
one, for it now seems almost certain that Paleohatteria is really
a Therapsid.
This suggestion was, in fact, really founded on such features as
the digital formula and the presetice of abdominal ribs, which
have proved to be common to all early reptiles. At the same
time, the structure of the neural part of the skull does recall
very strongly that of Sphenodon, or, so far as the interior of the
cranial cavity is concerned, a Tortoise.
As I believe very strongly in the value of this part of the
skeleton as an indicator of affinities, I think it probable that
Procolophon has some real relationship to the unknown group of
Cotylosaurs from which these two very diverse forms of reptiles
sprung. I have carefully examined the skeleton of Procolophon
in this connection, but can find no other resemblances between
it and the more modern reptiles which are not also common to
many other early forms. In particular the glenoid cavity seems
incapable of having given rise to that of reptiles with a single
coracoidal element, being curiously advanced in a different
direction.
For the opportunity of collecting the material on which this
paper is based, I am indebted to the assistance of the Percy
Sladen Trustees, and to G. W. Crozier, Esq., of Donnybrook,
W. Southey, Esq., of Tentergate, and P. Goosen, Esq., of Haslop
Hill, S. Africa, without whose hospitality and interest I could
have done nothing.
The beautiful preparation of the male skeleton was made by
Mr. R. Hall, of the British Museum.
A FOSSIL REPTILE.
EXPLANATION OF THE PLATES.
Puate I,
Procolophon trigoniceps.
Dorsal aspect of the nearly complete skeleton of a male (B.M. R. 4087).
Collected by the avthor at Haslop Hill, Dist. Tarka, 8. Africa.
Prater II.
Procolophon trigoniceps.
Ventral aspect of the specimen shown in PI. I.
Prats III.
Procolophon trigoniceps.
The specimen shown in Pi. I., from the left side.
Proc. Zoou. Soc.—1914, No. LI. 51
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MORMOSAURUS SEELEY.
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ON MAMMAL-LIKE REPTILES. 749
2. The Deinocephalia, an Order of Mammal-like Reptiles.
By D. M. 8. Watson, M.Sc., F.Z.8., Lecturer on
Vertebrate Paleontology in University College,
London. |
[Received May 7, 1914: Read May 19, 1914. |
(Plates IV., V.* & Text-figures 1-18.)
INDEX. Page
IGINGROGIECUNOS sedabo decoed soeBhe one coor once aacookee sen e OAT EC Racec EERE)
Systematic :
Mormosaurus seeleyi, gen. et SP. De ........cccccceeee vee ees 767
Pnigalion oweni, gen. et sp. n. WES Ee See TOT:
Lamiasaurus newtoni, gen. et sp. 1 Temes Ses SS ee ee erecey SOS,
Structure :
Tapinocephaloids :
Skull, statement of material ................. ......0..... 750
» Description ....... Gp Bone Ga Ra teecces soomepeny Goll
Post-cranial skeleton, material ........... Basic spree cha LO
S Description... .. 763
Dentition SRB SSE Hae ARB iano coe SERS EEAGc HU ONR ACER LER Een ERT ame LOT A
Titanosuchids :
SkllctabeMmentT Onn abe ria) ieee ees en 7/GO
6) IWESGRNAOM soo ccose Se stsnaasitects meee eed OO,
Post-cranial skeleton, material irae OAT ch Te 773
a ys IDESOMNUOD sciccocoobssascceesesocs 03
General discussion : :
Deuterosaurus a Tapinocephaloid ...... Bea seacrnone ane
Rhophalodon compared with Titanosuchids ............. 774
Affinities of Titanosuchus .. 774
Relation of Deinocephalia ‘to other South African
Therapsids ..... S Beak ded ender CAL
Relation of Deinocephalia to Pelyco cosauria . es Lo
The Auditory Arrangements of Therapsids ............... 779
ANG WON SATO coneacdoo.saqvsoceocséosoncn ecdeseaddbndacnoskannena eufhers
In 1851 Andrew Geddes Bain forwarded to the Geological
Society of London a series of fossil bones from the River Gamka,
Great Karroo, South Africa, which he recognised as different
from all the others he had collected.
These bones—a femur, humerus, ilium, vertebre, and portions
of skull-bones—were the first fragments of a Deinocephalian to be
found in South Africa. Many years before, however, the copper
mines of the Ural Mountains had yielded fragmentary bones
which were described by Kutorga; these are now known to
belong to Deinocephalians. The first South African type to be
described was TVapinocephalus atherstonei Owen, the snout of
which was excellently figured and shortly described in the
‘Catalogue of Fossil Reptiles of South Africa in the British
Museum. In this work Owen described some vertebre and
limb-bones belonging to Deinocephalia, but by a confusion in
localities attributed many of them to Pariaswurus. Some years
later Owen described the very fragmentary remains of Z%tano-
suchus.
* Wor explanation of the Plates see p. 786.
| 51*
750 MR. D. M. S. WATSON ON
The first account of a skull which was in any way complete
was written by H. G. Seeley on material of Rhophalodon and
Deuterosaurus from the Urals. Subsequently the same author
described a good skull belonging to the South African Museum
as Delphinognathus conocephalus.
Except for purely systematic descriptions by Broom of four
new genera and five new species founded on most unsatisfactory
material, nothing further was published till, in 1909, this author
gave a short and, as it has proved, somewhat inaccurate account
of two skulls in the British Museum collected by Prof. Seeley.
Subsequently Broom published a much better account of the
type skull of Delphinognathus and brief descriptions of several
new genera, one of which gave a complete knowledge of the
lower jaw.
Finally, Haughton has published a short account of a large
skull referred, perhaps correctly, to Vapinocephalus atherstoner.
The British Museum (Natural History) contains a large amount
of Deinocephalian material in the form of more or less frag-
mentary skulls and small associated sets of bones. From this it
is possible to get some idea of the form and structure of a
member of the group, and to give a nearly complete account of
the morphology of the skull.
The description of the Tapinocephaloid skull now given is
founded on the following material :—
I. R.1705. The type skull of Zapinocephalus atherstonei,
represented by the right side of the occipital region up to the
orbit, the left supraorbital region, and the anterior part of the
face.
Il. R. 3594. The skull already described by Broom as T'apino-
cephalus atherstonet which really belongs to a new genus (Pl. IV.
and text-figs. 1-4).
This skull has lost the basioccipital condyle and the left
quadrate region. It is very slightly distorted by pressure but
otherwise is very well preserved, having been completely freed
from a most intractable matrix by Mr. R. Hall, of the British
Museum. When Dr. Broom described it, the whole outer surface
was covered with a thin layer of matrix which has proved to be
readily removable, so that all the sutures of the face are clearly
and definitely shown; most of those figured by Broom are
wrong.
III. R. 3596 is a specimen in the Seeley collection from
De Cypher, Gouph, consisting of a skull (text-figs. 5-7) from
which the face in advance of the prefrontal had fallen away
before fossilisation, the two dentaries, with some limb-bones and
vertebre. Although also referred by Broom to Vapinocephalus,
this skull is generically distinct not only from this type but also
from R. 3594.
TV. 49385 is a specimen, Q of T. Bain’s collection, from Warm
Bad, Gouph, which consists of the posterior part of the skull,
MAMMAL-LIKE REPTILES, 751
lacking most of the quadrates, and the anterior part of the face,
which is only connected with it by documentary evidence (text-
figs. 8, 9). This specimen also represents a new genus.
Text-figure 1.
Pr. Fr. PR. FR.
Mormosaurus seeleyi, gen. et sp. n.
R. 3594 B.M.N.H. Type-skull. Right lateral aspect*, X +
J., Jugal; Lac., Lachrymal; Mx., Maxilla; Na., Nasal; P.O., Postorbital; Pr.
Pterygoid; Pr.FR., Postfrontal; Pr.FR., Prefrontal; P.Mx., Premaxilla;
Qu., Quadrate; Qu.J., Quadratojugal; S.Mx., Septomaxilla; Sq., Squa-
mosal; Tas., Tabulare.
Although these skulls belong to four different genera, they
agree so closely in their fundamental architecture that I have
used them all in the following description, which is, however,
founded as far as possible on R. 3594, from which the account
of the face and palate are almost wholly drawn.
[This method I hold to be perfectly legitimate ; it is as if we
were describing the characteristic structure of a mammal’s skull
from incomplete but complementary skulls of a bear, a dog, and
a cat.
ite Tapinocephaloid skull (text-figs. 1-9) has an extraordinarily
large and massive cranial portion beside which the often sharply
marked off face seems rather feeble. The whole of the bones
on the dorsal surface of the postorbital region fuse together and
thicken probably throughout the animal’s life, until as much as
%* Skulls from the Karroo of South Africa are usually distorted; in many cases
this is of the nature of a simple sheer, which is very easily corrected in making
drawings. In the figures of this paper this correction has been made, and each side
is restored by comparison with the other. Nothing is introduced into them without
clear evidence on at least one side. All the figures are projections and not perspective
drawings.
752 MR. D. M. S. WATSON ON
fifteen centimetres of solid finely cancellar bone may be formed
over the brain. The bones of the face, although they may be two
centimetres thick, never fuse, and were readily disarticulated in
the adult skull.
Text-figure 2.
IL [Pain
| si
UA \\ EEA
Pep Ex:
Mormosaurus seeleyi. Dorsal aspect of the type-skull, X 7.
Reference-letters as before, with :—Fr., Frontal; I.Par., Interparietal ;
Par., Parietal.
The whole occipital and otic region of the skull (text-figs. 3, 7)
has fused into a mass of bone in which no sutures are visible in
the material at my disposal. On the whole this region is very like
the corresponding part of Lystrosawrus which I have already
described. It forms a thick, vertically placed plate of bone
pierced through the middle by the relatively small brain-cavity.
This is throughout higher than wide, and has not been cleaned in
any specimens; fractures, however, show that it possessed the
characteristic Therapsid character of having the large opening to
the vestibule placed very low down in the skull. The floor of
the brain-cavity rises considerably towards the front.
The basioccipital condyle (text-figs. 5, 4, 7) is single, large, and
slightly pedunculate (R. 3596); it has a slight median depression
representing the notochordal pit. Below the condyle, the basi-
MAMMAL-LIKE REPTILES. 753
occipital forms a vertical area, which may be very high and wide
(text-fig. 7) (R. 3596) or comparatively small. The sides of this
area must be formed by the paroccipitals for some distance, as
the fenestre ovales lie at the sides well below the condyle. The
corresponding flat area in front is formed by the basisphenoid.
There are no distinct tubera basisphenoidalia, or basipterygoid
processes, but the posterior ends of the pterygoids articulate and
indeed fuse with the lower margin of the vertical area. From
Text-figure 3.
}
(Ce fi i ’ ip Wr SQ.
VS Borin Dy * Se
Mormosaurus seeleyi. Posterior view of the type-skull, x 4.
Reference-letters as before, with :—B.Oc., Basioccipital; Par.Oc., Paroccipital ;
P.T.F., Post-temporal fossa; Sr., Stapes.
between them the narrow parasphenoid rises as a vertical plate,
separated from much of the front of the basisphenoid by the deep,
short notch which is the pituitary fossa. This fossa is bounded
laterally by very low ridges which run upwards on the vertical
anterior face of the basisphenoid until they terminate in low pro-
cesses which are the processi anteriores prootici. The prootic of
course contributes to the fenestra ovalis, above which it is
perforated by the aqueeductus fallopii for the VIIth nerve.
The otic and supraoccipital regions of the skull are in contact
with the following bones :—
The upper outer corner of the posterior face of the large
paroccipital process with the tabulare.
The outer end and front face of the paroccipital process with the
squamosal.
The lower part of the front face of the paroccipital process with
the quadrate.
The posterior surface of the supraoccipital region is covered by .
the interparietal and tabulares. Its upper margin (and part of
the front face?) by the parietal (and postorbital 2).
754 MR. D. M. S. WATSON ON
The parasphenoid (text-fig. 9, Par.Sp.), as has been described
above, rises from the front face of the basisphenoid. It is very
short antero-posteriorly and rises nearly vertically in the skull
about to the level of the anterior inferior process of the prootic ;
its upper end is then split and receives the lower end of the
ethmoid.
The ethmoid is a remarkable bone extremely like that of
Endothiodon.
Its lower border is clasped by the split upper edge of the
parasphenoid, from which it rises as a narrow vertical plate; in
front this 1s continued to the roof of the skull, but behind it
Text-figure 4.
Mormosaurus seeleyi. Palatal aspect of the type-skull, x +.
Reference-letters as before, with :—Pr.V., Prevomer; Pat., Palatine ;
Tr., Transverse boue.
splits into two branches which rise to the frontals and leave
between them a small cavity for the olfactory lobes of the brain.
The nasal nerves leave by a pair of openings between the median
septal part of the ethmoid and the rest of that bone. The
MAMMAL-LIKE REPTILES. 755
ethmoid is received in a groove on the lower surface of the
frontals (and parietals?) and extends so far backwards that it
nearly, if not quite, touches the supraoccipital.
The interparietal (I. Pax.) is a flat bone entirely on the back of
the skull, where it overlaps the supraoccipital below and the tabu-
lares at the sides. The rest of its front face is in contact with
the hinder ends of the greatly thickened parietals.
The tabulare (TaB.) is a large bone forming a good deal of the
occipital surface. It is wedged in between the postorbital and
interparietal (R. 3594), and passes outwards above the post-
temporal fossa to touch the outer end of the paroccipital. The
outer part of its front face is in contact with the squamosal, and
its thick outer border joims with this bone to make a distinct
auditory meatus.
Text-figure 5.
r, 79>
ul!
Bk
te )-au.
Prigation oweni, gen. et sp. n.
Right lateral aspect of the type-skull. R. 3596 B.M.N.H. X }.
Reterence-letters as before.
The squamosal (S@.) is a large bone which has a powerful arti-
culation with the end of the paroccipital process; above the arti-
culation a strong ramus runs inwards along the front face of the
tabulare to overlap the postorbital. Immediately to the outside
of its articulation with the paroccipital process the bone overlaps
the upper end and some of the posterior surface of the quadrate.
Finally, the bone is completed by a powerful ramus which forms
part of the outer wall of the skull, articulating with the lateral
border of the quadrate and ending in suture with the quadrato-
jugal, jugal, and postorbital.
None of the specimens shows clearly all the sutures bounding
756 MR. D. M. S. WATSON ON
the parietal, but it is certain that the two bones met in a median
suture which is only interrupted by the long cylindrical tunnel
which forms the large pineal foramen.
The posterior border of the bone is covered by the interparietal.
Anteriorly it must meet the frontal, but the suture is never
visible. The lateral border is completely covered by the post-
orbital.
The excessive thickness of the bones makes them come into
contact with the upper border and anterior face of the supra-
occipital.
The postorbital (P.O.) is a large and very remarkable bone.
It forms the massive bar behind the orbit, and has a very long
suture with the squamosal in the zygomatic arch below the
temporal fossa.
Above the orbit in R. 3594 it meets the small post-frontal, but
in that specimen on its lower surface (on the right side) it can
be distinctly seen to join the prefrontal in a long suture running
inwards nearly to the middle line. The extremely thick and
massive bone then meets the parietal and covers the whole of its
outer surface, sending out a process in contact with the front face
of the interparietal and tabulare to be finally covered by the tip
of the parietal ramus of the squamosal, so that the temporal fossa
is entirely bounded by these two bones.
The sutures between the postorbital, frontal, and prefrontal
are not visible on the outside of the skull.
The postfrontal (Pr.FR.) is a very small bone forming a small
part of the orbital margin and entirely surrounded by the
prefrontal and postorbital (R. 3594).
The prefrontal (Pr.FR.) is a very massive bone forming the
front upper quadrant of the orbital margin. On its ventral
surface it has a long suture with the postorbital and likewise
joins the postfrontal, frontal, nasal, and lachrymal.
The lachrymal (Lac.) (R. 3594) is a small bone forming the
front of the orbit and articulating with the prefrontal, nasal,
maxilla, and jugal. There is no lachrymal foramen.
The nasal (Na.) (R. 3594) is a large bone which plays a great
part in the structure of the face. The two bones are in contact
in the middle line posteriorly, where they are very thick, but their
thinner anterior portions are separated by the very long internarial
processes of the premaxille. The outer border of the nasal has a
suture with the prefrontal, lachrymal, maxilla, and septomaxilla,
and is entirely excluded from the nostril by the latter bone.
The visceral surface of each nasal has a sharp ridge, the two
together forming a groove for the olfactory nerves.
The premaxilla (P.Mx.) of R. 3594 is a very remarkable bone.
It has a suture with its fellow, with which it forms the rounded
extremity of the snout. Its tooth-bearing margin is short and
wide, and its inner edge forms part of the anterior border of the
posterior naris. Medial to this opening it sends back a process
along the upper surface of the prevomer. The dorsal surface of
MAMMAL-LIKE REPTILES, 757
this process forms a floor to the anterior part of the nasal cavity,
and its outer border is in contact with the septomaxilla. The
outer side of the premaxilla has a powerful articulation with
the maxilla, In R. 3594 the facial part of the bone is continued
upwards by a long and slender internarial process which separates
the nasals for a great distance.
Text-figure 6.
Pnigalion oweni. Type-skull, dorsal aspect, X 2.
The maxilla (Mx.) is a bone with few remarkable features; in
front it joins the premaxilla, and thence backwards forms a
large area of the face. It is completely excluded from the border
of the external naris by the septomaxilla, with which in R. 3594 it
has a long suture. In this specimen it then has a short contact
with the nasal and is finally terminated by sutures with the
lachrymal and jugal. On the palate the maxilla is only repre-
sented by its wide tooth-bearing edge and forms part of the
border of the choana; for the rest of its length it is in contact
with the palatine and transverse bones.
The septomaxilla (S.Mx.) is a variable bone in Tapinocepha-
loids. In R. 3594 it has an internarial part which unites by
suture with the premaxilla to form a floor to the nasal cavity ;
it then sends back a large facial part which forms the whole of
the posterior border of the nostril and separates the nasal and
158 MR. D. M. S. WATSON ON
maxilla for a long distance; it is in contact with the maxilla
throughout its length, but is separated from the nasal just behind
the nostril by a foramen about a centimetre long and *5 cm. wide.
In 49385 the septomaxilla is entirely within the nostril, in
which it forms an irregular curved plate, having a suture with the
premaxilla and forming a rudimentary floor to the nasal cavity.
Tts outer border is in contact with the maxilla except for a small
foramen of irregular shape (text-figs. 1, 2).
The jugal (J.) is a comparatively small bone which forms about
a quarter of the orbital margin : in front 1t terminates in sutures
with the lachrymal and maxilla, and on its inner surface with the
transverse bone. Behind the orbit it has a suture with the
quadratojugal, squamosal, and postorbital.
The quadratojugal (Qu.J.) is a small bone on the side of the
skull; its end covers the outer border of the quadrate, from which
it is not separated by a foramen quadrati. Above it touches the
squamosal and in front the jugal.
Text-figure 7.
——
——
\\
NN
Pnigalion oweni. Type-skull, posterior aspect, X ¢-
Reference-letters as before.
The quadrate (Qu.) is a large bone whose lower border forms
the double condyle for the lower jaw; above this it rises in the
skull with its outer edge connected with the quadratojugal and
squamosal. The upper end of the bone and some of its hinder
surface is overlapped by the squamosal. .
Passing inwards from the body of the bone is a powerful
pterygoid ramus, which overlaps from the squamosal on to the
front face of the paroccipital process. Much of the rest of the
posterior face of this ramus is overlapped by the pterygoid.
Some distance above the condyle, where the pterygoid ramus joins
MAMMAL-LIKE REPTILES, 759
the body of the quadrate, there is a step which may (R. 3596) be
for the outer end of the extremely massive stapes.
The posterior end of the pterygoid is lightly applied to and
probably usually fused with the lower end of the basisphenoid at
the side of the parasphenoid. From the side of the bone in this
region (49385) a small process is given off which curves round so
as to shield the front of the fenestra ovalis. From this process a
slender rod rises very high in the skull with its posterior edge
nearly in contact with the front of the prootic: although not
separated by visible suture it is probable that this bone is the
epipterygoid. In R. 3594 there is a small fragment in contact
with the parietal in the region of the hinder end of the ethmoid
which may be the upper end of this bone.
From its articulation with the basiphenoid the pterygoid passes
forwards and outwards; the outer wing is the quadrate ramus,
which on account of the remarkable form of the skull and the
forward throw of the quadrate, forms a nearly flat, horizontally
placed sheet of bone overlapping the posterior surface of the
quadrate. In front of this the pterygoid narrows and then rather
suddenly widens and passes outwards to the transverse bone.
Much of the anterior border is in contact with the palatine in
a straight suture, but there is in 49385 a small medial process
which extends forwards to touch the prevomer.
The dorsal surface of the anterior part of the pterygoid is
produced into a high flange rising in contact with its fellow
(with which it may be fused, 49385) as a median septum in the
facial part of the skull.
The transverse bone (TR.) is separated from the pterygoid by
incomplete but quite definite and certain sutures on both sides
of R. 3594. It unites with the pterygoid to form a low flange
against the side of the lower jaw, and runs outwards, having a
long suture with the palatine in front to the maxilla and jugal.
Where the maxilla, palatine, and transverse meet there is
apparently a very small suborbital fossa only about °5 em. in
diameter.
The palatine (PAt.) is a bone of medium size which has a suture
with the transverse and pterygoid posteriorly, with the pterygoid
and prevomer medially; its anterior border forms a good deal of
the posterior naris and its outer edge has a long suture with the
maxilla.
The prevomer (PR.V.) is a rather remarkable bone. Behind it
has a suture with the palatine, for some distance its medial surface
is separated from that of its fellow by the thin vertical plate made
by the pterygoids. Further forward the two bones meet in the
middle line, and from this point forwards their upper surface isto a
variable extent overlapped by special processes of the premaxille.
The outer edge of the prevomer forms the whole of the inner
margin of the posterior naris. From the upper margin of the
bone a thin flange rises which leans inwards till its upper edge
touches that of the septum formed by the pterygoids. This flange
760 MR, D. M. 8S. WATSON ON
gradually declines in height, but in R. 3594 separates the sub-
narial process of the premaxille for some distance. _
In 49385 the prevomer underlies the premaxille to the
posterior edge of the alveoli.
Text-figure 8.
A
. * \ ‘ q \
AW"
ll .
os
/ J // y
1) Wn iY Cie
ZF ys
B
Lamiasaurus newtoni, gen. et sp. n.
A. Right lateral aspect of the type-skull, x 4. B.M.N.H. 49385.
. There is no evidence of the relative position of the back of the skull and the snout.
B. Posterior aspect of the type-skull, x 4.
C. Anterior part of the palate of the type-skull, <2.
MAMMAL-LIKE REPTILES. 761
The stapes (St.) in all Deinocephalia is a massive rod with few
features; in 49385 it apparently fuses with the posterior lip of
the fenestra ovalis. In R. 3596 it is of extraordinary size; the
distal end, which is directly and powerfully articulated with the
quadrate, being nearly 5 centimetres wide and 2 thick. It is not
perforated for a stapedial artery.
Text-figure 9.
——
=
ta
PAR
—— S
Type-skull of Lamiasaurus newtoni, external aspect of the bones of
the brain-case.
Reference-letters as before, with :—Eru., Ethmoid; E.Pr., Epipterygoid; Par.Sp.,
Parasphenoid; Prr., Pituitary fossa; VII., 1-foramen fallopii.
The Tapinocephaloid lower jaw is represented in the British
Museum only by the two dentaries of R. 3596 (text-fig. 16, A);
as Dr. Broom has recently published an account of the jaw this
lack is of little importance.
762 MR. D. M. S. WATSON ON
The post-cranial skeleton of Tapinocephaloids is represented by
the following material :—
The type of Tapinocephalus atherstonei :—The matrix of this
specimen isa dark blue grey limestone full of small lamellibranchs ;
these are the only bones in the collection in such a matrix.
The fragmentary skull, two nearly complete and several other
dorsal vertebree, and some caudals.
The type of Phocosawrus Seeley :—A sacrum and ten imperfect
dorsal vertebrae. The two ilia and pubes, one ischium, two im-
perfect femora, a tibia, two fragmentary scapule, and pre-
coracoids, one coracoid, two humeri, and one ulna.
The only fragment of skull associated with this is a bit of
palate showing part of the two pterygoids and a part of the
palatine. This agrees exactly with R. 3594, but is about half as
large again.
Text-figure 10.
- = iy }
A dorsal vertebra of the type-specimen of Tapinocephalus
atherstonei Owen, X}-
A. Anterior aspect. | B. Lateral aspect.
R. 1706 B.M.N.H.
Associated with the skull R. 3596 are several dorsal centra and
one arch, three femora, all extremely similar, a tibia, and a
humerus and atlantal intercentrum.
The atlas and axis of Moschops capensis have been described
and figured by Broom, Bull. American Mus. Nat. Hist. vol. xxxiii.
p. 139, fig. 5.
MAMMAL-LIKE REPTILES. 763
The anterior vertebre are not well represented in the Museum
collection, but have short deeply concave centra and massive
arches with short transverse processes ; the articular facet for the
rib seems to be continuous and to be carried on the arch and
Text-figure 11.
7 PINS
| GN
If}
f a
ha =
i
/
|
Nae
\\
Right scapula, coracoid and precoracoid of the type-specimen of Phocosaurus
megischion Seeley, X 3-
The part of the scapula between the lines which cross it is restored. It is known
from the evidence of other specimens that the length is approximately corvect.
C., Coracoid; P.C., Precoracoid; Sc., Scapula.
Proc. Zoou. Soc.—1914, No. LIT. 52
764 MR. D. M. S. WATSON ON
centrum equally (text-fig. 10). The later dorsals have centra like
those of the anterior vertebre, but the rib-facet is high up and
near the front border. The arches are high and massiveand carry
long transverse processes directed nearly horizontally. The zyga-
pophysial articulations are flat and inclined to one another and
the spines not very high.
The sacrum of Phocosaurus is composed of four vertebrae whose
centra are fused. The sacral ribs are articulated with both
centrum and arch; those of the anterior two vertebre are very
massive and of about the same size, the other two are much
smaller. One interesting feature is that the distal ends of the
anterior sacral ribs meet and fuse before they reach the ilium.
Text-figure 12.
Right humerus of the type of Pnigalion oweni, X +.
The caudal vertebre are very short, and the series belonging to
the type of Tapinocephalus seems to taper rapidly, so that the
tail was undoubtedly short.
The shoulder-girdle of Tapinocephaloids seems to be very large
and massive in comparison with the skull.
The scapula (Sc., text-fig. 11) is represented by its well-preserved
lower and upper ends in Phocosaurus and by an isolated bone
which gives the length. It has a flat blade with no indication of
an acromial process. The lower end has a long, straight face for
MAMMAL-LIKE REPTILES. 765
articulation with the precoracoid. Behind this it thickens and
forms the upper part of a glenoid cavity which looks outwards
and slightly backwards.
The precoracoid (P.C., text-fig. 11) is a very large bone arti-
culating by a long suture with the scapula and bearing a long
articular face for the coracoid. It is pierced by a foramen which
opens into a distinct pit on the visceral surface. The coracoid
(C., text-fig. 11) is a large bone which has a long surface for
contact with the precoracoid, and whose upper anterior corner
articulates with the scapula. Behind this contact it forms the
lower portion of the glenoid cavity, which is entirely formed by
the scapula and coracoid.
Text-figure 13.
Y <K (
NX
aN
‘ ys : \
BW Nana
|
Sacrum and pelvis of the type-specimen of Phocosaurus megischion Seeley ;
seen obliquely from front. + nat. size.
Iz., [lium ; Is., Ischium; Pu., Pubis; 8., Sacrum.
The humerus (text-fig. ]2) is a very massive bone with expanded
ends which are in planes at a considerable angle to one another.
The head is cylindrical and directly continuous with the enormous
deltoid crest, which, after extending more than halfway to the
distal end of the bone, suddenly subsides into the shaft. The
52*
766 MR. D. M. S. WATSON ON
shaft is relatively slender but very short, and soon expands into
the broad distal end. This is furnished with two articulating
faces: one almost entirely represented by a low knob on the
anterior face of the bone for the radius, the other on the end for
the sigmoidal fossa of the ulna. There is a large entepicondyle,
and a foramen piercing the shaft just at the end of the deltoid
crest.
The ulna is a short massive bone, with a pronounced olecranon
process and a facet for the head of the radius.
The pelvis of Phocosawrus (text-fig. 13) has already been de-
scribed by Seeley, and its general character will be best under-
stood from the figures of this paper. The interesting features
are the small pubic foramen, the absence of any obturator foramen,
and the enormous size of the acetabulum.
The four sacral ribs articulate with the inner surface by large
facets, which are continuous with one another.
Text-figure 14.
Left femur of the type of Prigalion oweni, X1. R. 3596.
The femur (text-fig. 14) is a very large bone with a rounded
head and long and large external trochanteric ridge, which slowly
subsides on to the shaft. The lower end is provided with two
condyles, which are only obscurely separated from one another.
The bone as a whole is flattened dorso-ventrally.
The tibia is a bone with very few features ; it has an expanded
head and a short, thick shaft.
MAMMAL-LIKE REPTILES. 767
The restoration of the skeleton (text-fig. 15) founded on all this
material gives one an idea, which is probably generally correct, of
the habit of the group. The striking features are the small head
and very massive limb-girdles. Phocosaurus is relatively more
heavily built than “ Pariasaurus” baine.
Text-figure 15.
Restoration of the skeleton of a Tapinocephaloid.
Skull, humerus, femur, and tibia from the type of Pnigalion oweni ;
other bones from the type of Phocosaurus.
Dentition and Systematics of Tapinocephaloids.
The type-specimen of TZapinocephalus atherstoner has been
ground down so as to show many sections of uncut tooth-crowns
and of the roots of functional teeth, Judging from these the
teeth are similar all round the upper jaw, there being no enlarged
canine.
The teeth-crowns, as shown by the series of sections which cut
them at many levels, consist of a sharp upstanding cusp, the outer
side of which is rounded and the inner side flattened ; from the
lower border of this cusp the crown is continued inwards on a
large shallowly concave area surrounded by ridges descending from
the sides of the anterior cusp ; these seem to be smooth, and the
outer is the larger. An isolated tooth (text-fig. 16, B) in the
Museum collection seems to agree with the tooth-crown restored
from the sections shown in this type-specimen.
R. 3594, which may be called
Mormosaurus seeleyi, gen. et. sp. n. (Pl. IV.),
has an extremely feeble dentition, which is quite uniform
throughout so far as can be judged from the usually imperfect
crowns and alveoli. Each tooth seems to consist solely of a cusp
which is oval in section and has a coarsely serrated edge.
R. 3596 may be called
Prigalion owent, gen. et sp. n.
The dentition of the upper jaw is not known, but there is no
sign of an enlarged canine in the dentary. The teeth seem to
be similar all round the mouth. Each tooth consists of a high
768 MR. D. M. S. WATSON ON
outer cusp with a rounded external surface and a deep groove
down its inner side. All round the inner side and front and
back of this cusp rises a strong cingulum, whose margin is raised
and serrated so as to form a cup in the middle of the crown.
Text-figure 16.
ii
ih
JN
if F Pl
Deinocephalian teeth.
A. Right dentary of type-specimen of Pnigalion oweni, X 5.
B. Tapinocephalus?, X 3.
C. Series of teeth of one individual of unknown genus, X 1.
No. 49585 may be called
Lamiasaurus newton, gen. et sp. n.
The dentition of the upper jaw consists of four large incisors
represented only by their alveoli, which are oval in shape and
MAMMAL-LIKE REPTILES. 769
suggest that the teeth may have been like those of Dewtero-
SMUTUS.
There is a single canine, which is large, strongly inclined
forwards, and circular in section. Behind this there are three
molars, shown in transverse section to be circular and relatively
small; it is probable that there was one more only.
One important difference between this type and the preceding
three is that whilst they all have very numerous replacing teeth,
which are not in the same stage and were thrust up indefinitely
like those of a crocodile, in Lamiasaurus there is no trace of
successional teeth.
The other types of South African Tapinocephaloids which have
been described are :—
Delphinognathus conocephalus, a much smaller form without
large canines, of which the details of the dentition are not
known.
Moschops capensis, also a small form with no canines and with
no details of the dentition.
Taurops macrodon, a large form apparently with teeth some-
what like those of Tapinocephalus, but not sufficiently described
or figured to be definitely identified at present.
Eccasaurus priscus, founded on a humerus of rather unusual
type, to which is referred a tooth of the general plan of Tapino-
cephalus but of a narrower oval form and more cuspidate.
Moschognathus whaitsi, founded on a figured lower jaw which,
as it seems to contain no teeth, is indeterminable.
The Titanosuchid branch of the Deinocephalia is represented by
the following skull-material in the British Museum (Natural
History) :—
An imperfect skull (Pl. V., & text-figs. 17, 18) consisting of
the dorsal surface from behind the nostrils to the back, with the
squamosals and part of the zygomatic arch; and a block which
has a rather poor fit with the upper part, but contains a good
deal of the maxilla and prevomers and a small fragment of the
palatine.
This specimen (R. 3595) was collected by Prof. Seeley at
Tamboer Fontein. The only other skull-materials I have used
are the premaxille of the type-specimen (49370), which itself
consists of the associated remains of two individuals of identically
the same size. The maxille of these agree exactly, so far as can
be seen, with that of the specimen mentioned above.
Nothing is known of the basicranial part of the skull, but the
supraoccipital above the post-temporal fossz is present in the
Tamboer specimen. It is a flat wide plate with rather large post-
temporal fossze bounding it below, and the upper part of the
deep and narrow brain-cavity impressed on its lower surface. Its
posterior surface is overlapped by the interparieta 1 and tabulares,
and its upper edge is in contact and perhaps fused with the
parietals.
770 MR. D. M. S. WATSON ON
A small part of the ethmoid remains in contact with the under
surface of the frontals, and is a small bone with a very small cavity
for the olfactory lobes of the brain.
The interparietal (I. Par., text-fig. 17) is a very large vertically
placed plate on the back of the skull. Its anterior face is in con-
tact with the upper part of the supraoccipital and the posterior
ends of the parietals. Laterally it overlaps the parietal rami of
the postorbital and squamosal and the tabulare.
Text-figure 17.
Titanosuchus. Dorsal aspect of skull. Premaxille from type.
The remainder from R. 3595 B.M.N.H. x2.
P.O., Postorbital; Sq., Squamosal; I.Par., Interparietal; Tas., Tabulare.
The tabulare (Tas., text-figs. 17 & 18 B) is a large bone,
whose medial border is covered by the interparietal and whose
MAMMAL-LIKE REPTILES, Hapah
anterior face has a contact with the supraoccipital, postorbital,
and squamosal. Its lower border forms the upper side of the
post-temporal fossa, and its outer edge is thickened and with the
squamosal forms a long, vertically placed auditory groove,
The squamosal (S8q., text-fig. 17) is only incompletely preserved ;
as shown it has a parietal ramus, which covers much of the front
face of the tabulare and extends inwards to overlap the posterior
end of the postorbital. With the tabulare it forms the auditory
groove on the postero-lateral corner of the skull, and beyond this
sends forwards a massive zygomatic part on the outer surface of
the skull.
Text-figure 18.
A. Titanosuchus. Wateral aspect of the same specimen as text-fig. 17.
B. Titanosuchus. Posterior aspect of skull. B.M.N.H. X ¢-
I.Par., Interparietal; Tas., Tabulare.
Except for the suture with its fellow and the frontal, the
parietal is completely shown. It is a small bone in contact with
its fellow except for the pineal foramen, which is a round hole.
The two bones form a special little projection raising the opening
more than a centimetre above the general line of the surrounding
bone.
(CU MR. D. M. 8S. WATSON ON
The sutures surrounding the postorbital (P.O.) are all visible,
but that with the postfrontal only on the ventral surface. The bone
forms a massive sheet at the back of the orbit, and forms the
hinder part of the extraordinary mass of bone which overhangs
it; it sends back a powerful process along the side of the parietal,
which forms the inner border of the temporal fossa, and thence
runs outwards along the front face of the tabulare to be covered
by the squamosal.
The sutures separating the postfrontal from the postorbital and
prefrontal are only visible on the under surface; they are straight
and run inwards for a very long distance with the postfrontal as
a narrow strip between them. It probably forms a good deal of
the knob above the orbit.
The sutures between the prefrontal and lachrymal ete. are not
visible, but on the under surface it seems that the nasals are
narrow bones, each carrying a sharp-edged rib, so as to form a
groove on the lower surface.
At the anterior end the specimen shows on the outer surface
traces of the sutures between the internarial processes of the pre-
maxillz and the nasals. It is evident that they were very long.
The premaxille known from the type-specimen have a flat facial
surface and rather broad internarial processes, the nares being on
the dorsal surface some distance behind the snout.
The palatal surface of the premaxilla is entirely formed by its
tooth-bearing edge, and the lateral border is in contact with the
maxilla.
The maxilla is a large bone with a great facial expansion.
Its palatal exposure is solely formed by its dentigerous margin, and
it is in contact with the palatine. Thereare large prevomers with
a long interchoanal bar, the dorsal surface of which rises into a
high thin flange. Of the quadrate only the articular edge is
known, which gives no indication of its size.
The type-specimen of 7itanosuchus ferox gives many bones of
the lower jaw, all disarticulated. The articular is a small bone
with a cotylus for the quadrate condyle, which apparently faces
very nearly backwards. The postarticular part is only repre-
sented by a small process on the lower side of the bone. The
anterior part rapidly narrows and finishes in a point. There
are well-marked articulating surfaces for the prearticular and
surangular, The angular is flat, and has a deep and very narrow
notch covered with a reflected lamina exactly as in a Gorgo-
nopsid. The dentary is a very massive bone, with a deep groove
on its inner surface for the prearticular, surangular, and angular
The bone seems to give evidence that there was no freely project-
ing coronoid process. One interesting feature is that there is a
very shallow, but quite definite, step in the dentigerous margin
behind the canine, as in Gorgonopsids, but, of course, relatively
very much smaller.
The lower surface of the dentary has a broad surface for the
splenial, which must have formed the whole of the lower border of
MAMMAL-LIKE REPTILES. WHS
the anterior part of the jaw, and had a large symphysis with its
fellow.
The post-cranial skeleton of Titanosuchids is represented by
two series of bones :—
The type-specimen of Zitanosuchus ferox (two individuals) :
neural arch and intercentrum of atlas, vertebree, scapule, cora-
coid, cleithra, clavicle, three humeri, femur, and fibula.
A series of associated bones of another genus, comprising
vertebre, ribs, ilium, humerus, femora, tibia, and skull-fragments.
The atlantal neural arch is absolutely identical with that of
the Dicynodont Aannemeyeria, as is the intercentrum. They
give evidence that the whole ‘complex was similar to that of
M oschops, as described by Broom.
The vertebre of 7itanosuchus are all extremely unsatisfactory,
but seem to be generally similar to those of Tapinocephaloids,
with longer neural spines and shorter transverse processes ; it is
just possible that the capitulum and tuberculum were connected
by a very thin web.
The other specimen has very high neural spines.
Certain fragments of bone belonging to the type of Titano-
suchus suggest the presence of relatively feeble abdominal ribs.
The scapule are very much weather ed, but are extremely large
and massive bones, giving no evidence of an acromion.
The coracoid is essentially identical with that of the Tapino-
cephaloids, and shows clearly that the precoracoid was excluded
from the glenoid cavity.
The cleithrum is a fine bone, nearly 8 centimetres wide and
2 thick; it has an articular facet for the front edge of the scapula.
What is almost certainly the lower end of the clavicle is
represented by an unsymmetrical sheet of bone about 20 em. by
30 em., which has the root of a powerful process arising from one
surface just within the edge. There is no other bone in the
skeleton it can possibly be, and, if it is correctly determined, the
interclavicle must have Reon an extraordinary wide flat sheet.
The humerus as figured by Seeley, Phil. Trans. 180 B, pl. xx
figs. 1 & 2, gives a rather misleading idea of the bone. Tt is in
essentials much like that of the Tapinocephaloids, but differs in
its relatively narrower lower end, in having the facet for the
head of the radius still more on the front of the bone, and the
two openings of the entepicondylar foramen on the same side of
the shaft. Finally, there is a small ectepicondylar foramen.
A single ulnare is present in the material; it is a rather small
but thick bone with a notch in one border, ‘forming part of the
ordinary foramen between the ulnare and the intermedium.
The ilium is very incomplete, but is a short bone, apparently
similar to that of the Tapinocephaloids.
The femur differs from that of the Tapinocephaloids in its
relative slenderness and the lesser size of the trochanter.
The tibia does not differ essentially from that of Tapino-
cephaloids.
U4 MR. D. M. S. WATSON ON
General Discussion.
That the animals whose structure I have discussed under the
heading of Tapinocephaloids are all closely allied admits of no
discussion, and is self-evident in every detail of their structure.
To the same group belongs the Russian type Dewterosaurus.
As shown clearly by Eichwald’s figures, the dentition is absolutely
characteristic, the powerful incisors which interlock are struc-
turally identical with those of Zapinecephalus. The enlarged
canine so clearly shown in his figures occurs in the remarkable
type Lamiasaurus which I have described above, and the presence
of only a single molar is a variation of slight importance. The
skull figured by Prof. Seeley has all the characteristic features
of the group, the broad zygomatic arch with the postorbital
obviously widely meeting the squamosal and the covering of the
side of the parietal by the postorbital (as shown and lettered
““post-frontal ” in Seeley’s figure) being identical with the struc-
tures in the South African forms. Even the slight difference in
the narrowness of the intertemporal bar and the erest on the
parietals can be matched in a fragment from South Africa in
the British Museum. Seeley’s figure also seems to show that
there was the very characteristic vertical area of basioccipital
below the condyle and the large quadrate characteristic of the
group.
The Russian type Rhophalodon and that described by Twelve-
trees as Cliorhizodon orenbergensis (R. 4077), which is perhaps
generically identical with the previously deseribed Deinosaurus,
are apparently closely allied to Vitanosuchus. Cliorhizodon (at any
rate, as shown by the beautifully preserved type-specimen in the
British Museum) is extraordinarily like a small Zitanosuchus,
as was, in fact, recognised in the original description. ‘There is
nothing in the whole structure of the skull to prohibit a
connection.
Whether Zitanosuchus is really closely allied to the Tapino-
cephaloids, or whether it is a specialised Gorgonopsid, as Broom
now believes, is perhaps more doubtful. It resembles the Tapino-
cephaloids in the extraordinary thickening of its skull and in the
whole texture of all the bones.
The whole structure of the temporal region, the form of the
squamosal, and the relations of the tabulares (postorbitals, parietals,
and squamosals) are identical in the two types. The very curious
auditory groove is the same in each type. The premaxille of
Trtanosuchus have a thoroughly Tapinocephaloid appearance, and
the position of the external nares far behind the end of the
snout on the dorsal surface is unparalleled in Therapsids, except
in that group.
If Rhophalodon is indeed related to Titanosuchus, then in the
large quadrate and the vertical area of basioccipital below the
condyle we have very striking resemblances to the contemporary
Tapinocephaloids.
MAMMAL-LIKE REPTILES, 775
Finally, so much of the palate of Vitanosuchus as is known is
quite unlike that of any Gorgonopsid, whilst it strikingly resembles
in detail that of Lamiasaurus. We are, I think, therefore
justified in regarding the two as divergent branches of the great
order Deinocephalia.
That the Deinocephalia are Therapsids has never been questioned
of recent years. The whole character of the limb-bones, par-
ticularly the humerus, is quite lke that of the corresponding
parts of an Anomodont, and is unknown in any other group.
The shoulder-girdle with the precoracoid excluded from the
glenoid cavity is extremely like that of a Gorgonopsid, and, except
for its lack of an acromion, that of Dicynodon. No similar
arrangement is known in any other group of reptiles. In the
more fundamental features of its skull-structure, the wide
occipital plate with the small and greatly separated post-temporal
fosse, the high brain-cavity, and the low position of the vestibule
of the ear, it agrees exactly with all South African Therapsids.
As is shown clearly by Dr. Broom’s figures of the lower jaw
and by that of 7itanosuchus, it has the characteristic flat angular
and notch of the Therapsids.
The group has, however, a very special importance, because alone
amongst the South African forms it retains a large quadrate.
The Tapinocephaloid quadrate is relatively as large as in any
other reptile—as large, for instance, as that of a Tortoise. The
importance of this is that it shows that the reduction of the
quadrate, which is so noticeable a feature of the Anomodonts
and “‘Carnivorous” Therapsids, is not an essential feature of
Therapsid structure, but has developed in comparatively ‘late
times in at least two branches independently (I intend to adduce
evidence in support of this statement in a later paper). Some
years ago, in describing the skull of Diademodon, I pointed out
that this reduction of the quadrate was part and parcel of a whole
series of changes which led to the reduction of all regions of the
skull (the basisphenoidal, basioccipital, and exoccipital regions
in particular) which lay below the base of the brain. This
generalisation is justified by all subsequent work on Therapsids,
and is well illustrated by the great depth of the basicranium in
Deinocephalia, an arrangement which may to some extent be
paralleled in Deinosaurs.
The exact relation of the Deinocephalia to other South African
groups of Therapsids is not easily determined.
The skull of Zitanosuchus bears a very considerable resem-
blance to that of a Gorgonopsid, modified, of course, by its great
thickness and the large bony bosses over the orbits. It is dis-
tinctly more primitive in many features—for instance, in the less
widely spread zygomatic arches. The skulls of the more primi-
tive Gorgonopsids, however, show no trace of the vertical area
of basioccipital below the condyle; and there are no features of
the auditory region of the brain-case which in any degree suggest
that they have been derived from a type with a Deinocephalian
776 MR. D. M. 8. WATSON ON
structure. It will be remembered that Rhophalodon, which is in
many ways extremely like a Gorgonopsid, has this characteristic
Deinocephalian structure. So far as the lower jaw goes, the
Gorgonopsids could be derived directly from a lightly built
Titanosuchid. The post-cranial skeleton, so far as is known, is
also very similar, allowing for the differences due to the great
weight of all well-known Deinocephalians or, using as a term of
comparison, the relatively slender bones from the Ural copper-
mines.
In fact, it is legitimate to assume that Deinocephalia and the
Gorgonopsids arose from a not very distant common ancestor, but
that they subsequently pursued quite different paths.
Perhaps the most interesting comparison is with the Anomo-
donts. In many features (the upturned parasphenoid and the
structure of the ethmoid, for example) there is a distinct resem-
blance between the two groups, which also resemble one another
to some extent in the shortness of the brain-cavity—in some types,
at any rate. Further, many Anomodonts have a considerable
vertically placed area of bone below the basioccipital condyle,
which may be compared with that in Tapinocephaloids. As this
area does not exist in Yndothiodon, whilst it is most pronounced
in Lystrosaurus, it is probably a secondary specialisation inde-
pendently acquired within the group. Connected with this
feature, however, is the development of that curious process of
the vestibule which carries the fenestra ovalis down to the lower
border of the skull. This curious detail is found even in
Endothiodon, a rather primitive type, and is apparently quite a
fundamental character of the group.
Although the cavity of the inner ear has not been cleared in
any Deinocephalian, there can be no doubt that a similar structure
did occur in that group—a fact which suggests a nearer relation-
ship between the two groups than either of them hold with the
Gorgonopsid line.
At the same time the Deinocephalia differ from all other
South African Therapsids in the mode of articulation of the ribs
in the dorsal region.
The most interesting comparison is between the Deinocephalia
and the Pelycosauria, using that term to cover all the “Texas ”
Therapsids.
If we compare a Deinocephalian with the very primitive
Varanosaurus, we find many resemblances. We have, to begin
with, the characteristic Therapsid characters of the occiput and
lower jaw. In addition, we find that there are striking resem-
blances between the quadrates of the two types in their size,
relation to the squamosal and quadratojugal, and in the power-
ful step for the distal end of the stapes. Another resemblance is in
the form of the squamosal, which in both types is a simple sheet
folded round the back of the quadrate, and not produced outwards
as in the Anomodonts and Therocephalia. The occurrence of a
MAMMAL-LIKE REPLILES. ata
vestigial ‘ supratemporal” in Varanosaurus is a primitive feature
of little importance.
Comparison of a Tapinocephaloid with Dimetrodow leads to
some interesting results, as Broom has already pointed out.
There is a resemblance in shape, owing to the very short teim-
poral regions and high and compressed form of the skull in
both groups. The basicranial region of Dimetrodon, although
more specialised than that of Varanosaurus, shows no suggestions
of the characteristic Deinocephalian structure, and is, in fact,
extremely like that of the more primitive of the Gorgonopsids.
The very peculiar parasphenoid is identical in the Deinocephalia
and Dimetrodon, and it is probable that the ethmoids are also
similar. The quadrate of Dimetrodon is modified from its
primitive condition by that development backwards of the
posterior angles of the skull which is shown by the curious
backwardly directed processes of the outer ends of the paroccipital
processes. Making allowance for this modification, its relations to
the surrounding bones are essentially those held by the corre-
sponding bone of Mormosaurus, so far, at any rate, as can be
judged from published descriptions. In the palate also there are
distinct resemblances in the development of the high vertical
flanges from the pterygoids and prevomers. The two types are,
however, specialised in directly opposite directions, one with a
short face for a herbivorous diet, the other with the elongated
gape which is necessary to a carnivorous animal with the charac-
teristic reptilian habit of grasping its prey after a single snap.
The occiput of Hdaphosaurus, as figured by Williston and Case,
is extremely like that of Pnigalion, and, if we may judge by
Varanosaurus, was essentially similar in structure.
Finally, as Broom has pointed out, the face of Dimetrodon is
structurally very similar to that of such a Deinocephalian as
Mormosaurus—it is perhaps even more similar to that of Deutero-
SaUrUs.
On the whole, the Deinocephalian skull resembles that of the
Pelycosaur more closely than it does any other South African
Therapsid. |
As Broom has already shown, the lower jaw of a Tapino-
cephaloid, except for the differences due to the oppositely
specialised dentitions, is structurally very similar to that of
Dimetrodon. It differs, however, in the more greatly developed
Therapsid notch.
Broom has already shown the very striking resemblance between
the Pelycosaur vertebrae (particularly those of Dimetrodon) and
those of Tapinocephaloids.
In the rest of the post-cranial skeleton, however, there is very
much more similarity between the Tapinocephaloids and the
other South African Therapsids than between that group and
the earlier Pelycosaurs.
In the cartilaginous shoulder-girdle the precoracoid is entirely
778 MR. D. M. S. WATSON ON
‘excluded from the glenoid cavity, and there is no trace of the
screw-shaped form of the articular surface which is an essential
feature of the primitive type. The humerus also is of a more
modernised form, its action being an up and down movement
instead of one which is nearly parallel to the ground. ‘The
very broad lower end of the clavicle of Titanosuchus is, however,
a point of resemblance with the earlier forms.
In the pelvic limb we find a somewhat similar mixture of
characters.
The whole pelvis, for example, is somewhat similar to that of
Dimetrodon, particularly in the production of the outer corner of
the front border of the pubis and its deflection, more so perhaps
than to any South African type; but the femur is of a modernised
type strongly resembling that of Dicynodon.
The final result is that the presence of the Deinocephalia
makes it impossible to exclude the American Lower Permian
and Carboniferous Pelycosaurs from the later South African
Therapsids. Such a division could at any time have been drawn
only on the more primitive limbs and large quadrate of the early
forms; the fact that in Deinocephalia we have types with a
quadrate as large as that of the Pelycosauria combined with
modernised limbs renders the foundation of a great group-
division on these characters quite impossible.
For this great stem ef the Reptilia, including all the mammal-
like reptiles, many names are available. I am myself inclined to
extend Broom’s Therapsida, a most appropriate name, to the
whole of them, but I fully recognise that Cope’s earlier names of
Theromorpha and Theromora have been used in the same sense ;
these names were never very clearly defined by Cope, and have
at one time or another included nearly all Permian reptiles. If
anyone should wish to resuscitate these names in this connection,
I would point out to them that Owen’s term Anomodontia was
used by that author in 1860 in a wide sense to include the
Dicynodonts and also carnivorous Therapsids from South Africa,
and has at least as good a claim to be used as Cope’s later terms.
Discussion of Special Features of the Skull.
The septomaxilla of Deinocephalia is of interest on account of
its variability. It resembles that of all other Therapsids, except
certain Cynodonts and Anomodonts, in the fact that there is a
foramen behind it opening from the outer surface into the nasal
cavity. It is of interest that this foramen was first described by
Case in Dimetrodon. It is evident in the majority of the more
primitive South African “Carnivorous” Therapsids, and is clearly
shown in a British Museum skull of Hndothiodon and in
Prof. Sollas’s model of Dicynodon. Its function is unknown, the
only suggestion I have yet found is that it may mean that the
ductus naso-lachrymalis opened on the surface; it will be re-
membered that in the small Temnospondylous Stegocephalian
MAMMAL-LIKE REPTILES. 779
Micropholis this duct runs forwards completely in the lachrymal
bone to the posterior end of the septomaxilla, and it appears from
its mode of development to have been primitively a mere surface-
groove.
The squamosal of Deinocephalia is of some interest; in its
relations to all other bones, particularly those of the otic capsule,
the quadrate, tabulare, postorbital, and parietal, it agrees exactly
with the corresponding bone of the more primitive Gorgonopsids,
which is clearly identical with the squamosal of Diademodon,
which is clearly homologous with the bone of the same name in
Mammals. The squamosal of a Deinocephalian is clearly the
same as the large temporal element in Varanosaurus, having
similar relations to the paroccipital, postorbital and tabulare, and
only differing in retaining its primitive connection with the
pterygoid. As this type retains a small ‘‘supratemporal,” there
is now no doubt that the squamosal is the outer of the three
temporal bones in primitive Reptilia.
The Auditory Arrangements of the Therapsids.
Of recent years belief in the truth of Reichert’s hypothesis of
the homologies of the mammalian ossicula auditus has been
gradually growing, and the fine work of Gaupp has practically
placed it beyond question. At the same time, Broom, following a
suggestion of Seeley’s, has shown how in the gradual increase in
size in the dentary and the decrease of the other elements of the
jaw and the quadrate, the “carnivorous” Therapsids gradually
approach the mammalian condition. The same author has pointed
out that the stapes is articulated with the quadrate in Therapsids
just as is the stapes of mammals with the incus. Finally,
following on my own more accurate account of the Therapsid
jaw, Mr. Palmer has shown the extraordinary similarity in detail
between the lower jaw (including the dentary, malleus, pre-
articular, and tympanic) of a mammary foetus of Perameles and
that of the advanced Cynodont Diademodon.
The homologies of the various bones being now placed beyond
dispute by the work of these authors, it is desirable to carry the
inquiry further and discuss the position and changes of the
tympanic membrane and other parts of the ear.
The stapes of Therapsids is in all cases where the condition is
known articulated with the quadrate. In the following types,
Cynognathus, Trirachodon, Nythosaurus, Arctognathus, Lyco-
saurus, Dicynodon, Lystrosaurus, Dicelurodon, Kannemeyeria,
Mormosaurus, Prigalion, Lamiasaurus, Dimetrodon, Varanosaurus,
which cover the superorder very fairly, the stapes has actually
been seen in place; the facet on the quadrate for its distal end is
shown in very many more forms.
Quite recently, when discussing the primitive Therapsid Varano-
saurus, | pointed out that in many ways it greatly recalled the
Proc. Zoou. Soc.—1914, No, LILI. 53
780 MR. D. M. S. WATSON ON
Cotylosaurian family of the Captorhinide, so much so that a
blood-relationship between them seemed most probable.
Prof. Williston had still earlier expressed a similar opinion :—
“That we have in Labidosaurus and its allies a persistence of
those generalized characters which gave origin to the peculiar
specializations of the Pelycosaurs.” One of the peculiarities of
these Captorhinids, in which they differ from all other known
Cotylosaurs, is the fact that they have a stapes whose distal end
is articulated with the quadrate. We have therefore some
justification for assuming that this peculiar condition in the
Therapsids has not arisen within that group, but has descended
to it from its ancestors.
I have shown that there are very strong reasons for believing
that the reptiles have arisen from the embclomerous Stegocephalia,
which are the only group of Tetrapods known in Lower Carboni-
ferous rocks. In figure A, in the plate of my paper on ‘The
larger Coal-Measure Amphibia,” Manchester Memoirs, vol. lvii.
pt. i., there is shown a very definite depression on the upper
surface of the quadrate and pterygoid. ‘This definite pit occurs
in all embolomerous Stegocephalia in which I have examined
this region, and is obviously of importance. The only explanation
I can find for it is that it received the outer end of the stapes.
It is important to note that in the uncrushed skull it must have
looked towards the auditory region. There is thus a suggestion
of evidence that the Cotylosaurs of the family which gave rise to
the Therapsids and the Captorhinids received the condition of
having the distal end of the stapes articulated with the quadrate
from their amphibian ancestors, as they in turn passed it on to
their descendauts.
There are strong reasons for believing that the Stegocephalia
have arisen from the Rhipidistian group of the Crossopterygian
fishes (the Osteolepids), which were no doubt technically, though
probably only to a slight extent, functionally hyostylic. In these
fishes the hyomandibular, which has been seen only by Dr. Traquair
in Rhizodopsis (an observation I have been unable to check from
the material in the British and Manchester Museums, which
includes at least the greater part of that which was before
Dr. Traquair), no doubt articulated with the otic region of the
cranium and with the quadrate, as in all fishes. There are many
reasons for believing that the stapes of a Tetrapod is homologous
with the hyomandibular of a fish, and it is probable that in the
embolomerous Stegocephalia, which had just arisen from fishes, the
primitive connection between the distal end of the hyomandibular
or stapes and the quadrate was retained. In later Amphibia
(Eryops, Trimerorachis, Cyclotosaurus, to mention only types in
which it is known in place) it lost this connection, and its distal
end is connected with a tympanic membrane stretched across the
otic notch.
It thus seems to be fairly probable that the connection between
the distal end of the stapes and the incus in a mammal has been
MAMMAL-LIKE REPTILES, 781
directly derived from the connection of the distal end of the
hyomandibular and the quadrate in its far-off fish-ancestor, and
that this connection has never been lost in the phylogeny of the
group.
The most primitive lower jaw known amongst Therapsids, that
of Dimetrodon, as 1t has been described and figured by Case,
Williston, and Broom, is differentiated from that of a member of
any other Reptilian supererder by the characters of its posterior
part, and particularly the angular. In Dimetrodon the angular
is essentially a flat plate whose upper border overlaps the sur-
angular and is itself overlapped by the dentary in the usual way,
whilst its lower border forms the lower border of the jaw. The
posterior end of this bone is separated from the outer side of the
prearticular by a notch, so that the border stands out freely.
The whole jaw is very narrow from side to side, the Meckelian
vacuity being reduced to a very narrew slit between the pre-
articular and the surangular. The flat plate-like angular and
the notch in its posterior border are the characteristic features of
this jaw, and are found in all Therapsids which are properly
known and in no other reptiles whatsoever.
The Deinocephalian jaw only differs from that of Dimetrodon
in the further enlargement of the notch, which in this type
extends forwards so as to form a deep pocket in the substance of
the angular. In such Therocephalians as Seylacosaurus the con-
dition of the angular is almost exactly as in Deinocephalia, there
being a deep but very narrow pocket in the substance of the
angular which forms a large reflected lamina outside it. The
foramen shown in the angular in many of Broom’s figures of
Therocephalian skulls has no existence in fact, as such, but is
merely the anterior end of the notch.
In the Gorgonopsids there is the usual notch and reflected
lamina, which is, however, small, never reaching the size of that
of Scylacosaurus. This notch in the higher Gorgonopsids, e. g.
Arctognathus, seems to move relatively further forward. Finally,
in the Cynognathids, which have a “‘ look” of the Gorgonopsids,
of a character that cannot be intelligibly expressed in words but
is very striking to anyone accustomed to handling the two types,
and suggests that they are in all probability genetically related,
the notch has moved still further forward, and the reilected
lamina now stands downwards from the rest of the bone as a
very slender process. The extraordinary resemblance of this
bone to the tympanic of the pouch feetus of Perameles (or of
Dasyurus) seems to me to raise a very strong case for believing
that Cynognathus and its allies had a tympanic membrane spread
between the divergent branches of its angular. ‘The actual shape
of the bone makes such a, position possible, the hinder end of the
membrane being carried by the ridge on the lower side of the
squamosal, which lies just outside the end of the paroccipital
process, and on the musculus depressor mandibuli, if such a
muscle be present. A membrane in this position stands nearly
53*
782 ; MR. D. M. S. WATSON ON
vertically in the skull, and les at the lower and inner end of the
groove which Broom and ‘I, following the original identification
of W. K. Gregory, regard as an external auditory meatus. If
this position was really that held by the tympanic membrane in
Diademodon, then in that type the tympano-tubal cavity had
already grown up round the bones of the back of the lower jaw.
{f this position of the membrane be correct, by tracing back
along the series of Gorgonopsids to the Deinocephalia. we ought
to have self-consistent results throughout, and to arrive at a
condition in primitive forms which is not inconsistent with that
found in known Reptilia. The series of Gorgonopsids at my
disposal gives a sufticiently close morphological series to show
with certainty that :——
lst. The noteh in the hinder border of the angular of a Deino-
cephalian is homologous with the wide triangular notch
in Diademodon.
Ind. That the very small but distinct auditory groove described
above in Deinocephalia is homologous with that of Dia-
demodon.
3rd. That the ridge to the outer side of the paroccipital on the
squamosal in the two types is the same.
These homologies lead us to expect that the tympanic membrane
of Deinocephalia should be attached to the squamosal just outside
the paroccipital process, to the posterior surface of the quadrate,
and to the edge of the reflected lamina of the angular. I believe
such an attachment to be workable inasmuch as a tympanic
membrane attached to it might be flat.
The groove for the external auditory meatus in Deinocephalia
is of great interest. It is undoubtedly homologous with that of
Diademodon, and I think almost as certainly with that of a
mammal. It gives clear evidence that even at this time the
tympanic membrane (whatever its precise situation and attach-
ments) had sunk in and was no longer directly exposed on the
outside of the head. It 1s formed by a groove between the outer
edge of the tabulare and the squamosal; this groove terminates
suddenly where the tabulare ends in a definite projection from
the back of the skull which must mark the upper insertion of
the membrane.
It is known from the evidence of specimens which show the
complete stapes in position—Hryops, Trimerorachis, Cyclotosaurus,
ete.—that in Stegocephalia the tympanic membrane was stretched
between the tabulare and the squamosal across the otic notch
on the upper surface of the skull. By direct tracing back of
homologous parts we have seen that in Deinocephalia the upper
end of the membrane should be attached to the squamosal and
the extreme distal end of the tabulare: that is, in the exact
position we know it to have been attached in Stegocephalia
related, although remotely, to the Deinocephalian’s amphibian
ancestors.
MAMMAL-LIKE REPTILES. 783
It will be convenient here to leave this particular line of
argument, and starting from the normal conditions of the mem-
brana tympani in Reptiles, try to reconstruct the arrangement in
Dimetrodon, and so work back to the termination of our original
argument at the Deinocephalia.
In Lizards the insertion of the tympanic membrane is on to
the back of the quadrate, the squamosal just outside the par-
occipital process, and the retro-articular portion of the lower jaw.
In Pelycosaurs and other Therapsids the retro-articular portion
of the lower jaw is of insignificant proportions, at its largest in
Dimetrodon. In Deinocephaha and Dicynodon it is represented
solely by a small process from the articular directed as much down-
wards as backwards. The definite presence of this process in these
more primitive forms and its still further reduction in later types,
seem to show that some Therapsid ancestor had a relatively
large retro-articular process. As in all reptiles which have such
a process its outer side is covered by the angular, we may assume
that in this hypothetical ancestral Therapsid the retro-articular
portion of the angular played some part in the support of the
tympanic membrane. If now in such an animal we make the
tubo-tympanal cavity grow forwards and the membrane to keep
pace with it in such change, we must also move forward the upper
edge of the retro-articular part of the angular to which the
membrane is attached. As it is essential to keep the stretched
membrane clear of other bones, and the articular and quadrate
are by hypothesis fixed, we can only do so by separating the edge
of the retro-articular part of the angular from other bones and
moving it slightly outwards: this on the theory which I am at
present expounding is the origin of the Therapsid notch, the
upper edge of the reflected lamina being phylogenetically the
upper border of the retro-articular portion of the angular, and
retaining that connection with the lower edge of the membrana
tympani which occurs in Lizards. By this method we arrive at
a, position of the tympanic membrane in Deinocephalia identical
with that deduced by tracing the conditions down from Cyno-
gnathids, the arrangement in which was determined by assuming
that the great resemblance between the angular of these types,
with their long, slender, downwardly directed “ reflected lamina,”
and the tympanic of a pouch-young of Perameles with its slender
lower limb, was a real one. The fact that it is possible to trace a
sequence of stages with only very few hypothetical intermediates
between the actual condition of the membrana tympani in lizards
and that in embryo mammals, and that these stages are in the
correct time order, and are each self-consistent, seems to me to
establish a probability that the assumption on which they depend,
i. e. that the reflected lamina of the Therapsid angular carried the
tympanic membrane, is a justifiable one.
T am quite aware that the shape of the edge of this lamina in
some types, as for example in Hndothiodon and Anomodonts
generally, is inconsistent with the view that it carried the
.
784 MR. D. M. S. WATSON ON
membrane, but as in all these types the squamosal never shows
any suggestion of an auditory groove or of the attachment of a
membrane, I think it possible that they were without one, a not
unknown condition.
In 1910 W. K. Gregory, from the position of the anditory
groove in Cynognathus, inferred that the tympanic cavity and
membrane were below the reduced quadrate and articular. From
these relations and the fact that in the ontogeny of a mammal
the tubo-tympanal cavity grows up round the auditory ossicles
which arise outside it, he suggested that phylogenetically this
upgrowing of the tubo-tympanal sac around the vestigial quadrate
and articulare may have caused them to share in its vibrations,
and thus to take on an incipient auditory function before their
old suspensory function had ceased.
This is exactly the conclusion to which the above discussion,
founded mainly on quite different evidence, has led us.
In mammals the centre of the tympanic membrane is placed
in connection with the chain of auditory ossicles by the handle
of the malleus being fastened to its middle layer. In no
Therapsid that I have yet examined is there a process of the
articulare which could touch the membrane, so that it Is m-
herently probable that the manubrium is a mammalian innovation.
Tn ontogeny it arises very late, chondrifying much later than the
incus and body of the malleus in erameles, but being apparently
a veal part of the latter bone.
It thus seems impossible that the membrane of Therapsids
should be brought into connection with the fenestra ovalis in the
ordinary mammalian way, and as the whole of the preceding
arguments are meaningless if they have no membrane, it seems
certain that another connection between the stapes and the
membrane must have existed.
In a remarkably able and suggestive paper, Dr. Gregory has
put forward the following explanation :—
“That in the most primitive Cynodonts, such as Bauria, there
was an extra-columelia, resting against a tympanic membrane
behind the squamosal, which had been differentiated out of the
tissue lying between the endodermal epithelium of the tympanic
eavity and the epidermis: that with the spread of the tympanic
cavity the differentiation of the future tympanic membrane also
spread, until it included the stretched skin on the posterior end
of the jaw below the quadrate and articular and above the
angular: that concomitantly with the reduction of the quadrate
and articular and the detachment of the angular and goniale
from the dentary, the newly differentiated portion of the tym-
panic membrane became functionally more active than the old
‘reptilian’ portion : that in this way the old membrane together
with the extra-columella became vestigial, while the new membrane
beeame altogether free from the dentary, but remained fastened
both to the angular, which gave rise to the tympanic bone, and
to the retro-articular process of the articular, which gave rise te
MAMMAL-LIKE REPTILES. 785
the manubrium of the malleus. With the reduction of the
‘reptilian’ tympanic membrane the hyoid became separated from
the extra-columella (as it does in many lizards) and migrated to
a new insertion on the periotic.”
The foregoing discussion will have shown how much of truth
there may be in this brilliant hypothesis of Gregory’s. The
suggestion that the stapes of Therapsids although directly arti-
culated with the quadrate may have been connected with the
membrane by an extra-columella, is exactly paralleled by the
actual conditions in Mosasaurs, where the slender stapes is in-
serted into a deep pit on the ner side of the quadrate, but is
connected with the tympanic membrane by a process passing
through the large special notch on the back of that bone. This
process like the membrane is sometimes strongly calcified. The
whole arrangement is after all only a further development of
conditions commonly found in lizards.
The facts as we know them in Diademodon, or rather the in-
terpretations of those facts offered above, show that the reptilian
portion of the tympanic membrane had actually been reduced to
very small dimensions by a steady process connected with and
no doubt induced by the same factor as the degeneration of the
quadrate and the thinning of the basisphenoid and basioccipital.
The facts also seem to show that the manubrinm mallei is
altogether a new formation, developed whilst the articular was
losing its suspensory function.
If the explanation of the meaning of the observed series of
changes in the articular region of the skull of Therapsids which
has been presented above be true, or even if it contains but a
small element of truth, it presents us with a good illustration of
the fact, to my mind patent in the development of every organ
of the body which is known, that the Therapsids from the moment
of their initiation were committed to the final development of
a mammalian structure. Different branches of them proceeded
with their modifications to different degrees, and at very different
speeds, but so far as the evidence goes, and it is no more than
suggestive at best, their evolutionary change, in fundamental
features, were always directed towards the final development of a
mammalhan structure.
I wish to acknowledge my indebtedness to the Percy Sladen
Trustees, who assisted me in visiting the South African Museums.
To Dr. Smith Woodward IL owe the opportunity of describing the
material in the British Museum (Natural History), which is the
basis of this paper, and I also owe to him and to Dr. C. W.
Andrews thanks for their many kindnesses during my work at
the Museum. Finally, I have to thank Mr. R. Hall, chief
‘“mason” at the Museum, who developed the skulls described in
this paper from a matrix which in many places is as hard as
flint.
l on Pet a { ae y
es) me ‘i ae epee ee ey
$BGx i a ene KE REPTILES. ee
. } : \ : fo) ~1 : i;
| a eS
EXPLANATION OF THE PLATES. = 2 | |
Prats IV. wae 3 ee
Fig.1. Skull of Mormosaurus seeleyi, gen. et sp. un. Type specimen (R. 3594).
From side. About + nat. size. We
Fig. 2, Skull of Mormosaurus seeleyi, gen. et sp. n. Type specimen (R. 3594). —
Obliquely from front. About } nat. size. at
Puate V.
- Skull of Titanosuchus feror Owen. (R.3595,) From above.
About 3 nat. size.
PZ.S.1914.PRESTON.PLI.
2x.
eed a,
eC GC oxPe. a3xea.
SEALE
G.MWoodward del.et lith. Huthimp.
NEW ZONITIDA FROM EQUATORIAL AFRICA.
PAS. Lo] PAE See
LO xl.
es
Bey ale,
G.M.Woodward del.et lth. Huth imp.
NEW ZONITIDA FROM EQUATORIAL AFRICA.
P2Z.5.1914 PRES YON. Pian
ISeae
G.M.Woodward del.et lith, Huth imp.
NEW ZONITIDA FROM EQUATORIAL AFRICA,
ON NEW GENERA AND SPECIES OF ZONIVIDA FROM AFRICA. 787
43. Diagnoses of new Genera and Species of Zonitidee from
Equatorial Africa. By H. B. Preston, F.Z.8.
[Received April 3, 1914: Read May 5, 1914. ]
(Plates I-III.*)
INDEX. Page
Geoomap incall apeeses ithe penetra cone oa eictade cota ON
Systematic :
IOV GO DANG D1 il Scqubdodsoce dco teen sas ebIaagEae Om a)
IE aPURT AMG FOE NG goo bscee dsocns ecocaacen or crgeee, UL
TIGRE ANG, (RDBlo We do 30 dclachashonwedbeone can ehearen WEL)
J GERDA Gi, (RDN Whe, csp baboceacesabaanaqaucoceepecesecen | tll)
INCU T RA iy ERO Wo shnecsecacnsecannboeeseeeoneenss telUY4
DGIRETC AH Kei E295 We esnos nbudeoedaHeascodespaaace sencdd mae.
WRGLERSANG, BOUS Silo enacbsaqnon copaosossoncensancee | AUB)
IRON AL NING sodees beg seasacmes ooo cd=soc dapeepere yO
Faltloonella, gen. MN. ....-..-.. eee eceeeeces. 1009,
And 74sp.n., 1 sub-sp. n., and 2 var. n.
In working at the present family the author has found it
necessary to refer a number of the species to new genera, having
found that the Zonitide of the Central African region have been,
hitherto, in common with those of many other geographical areas,
massed together in a few genera, so that these now contain many
widely divergent forms; in very few instances has he found, as
might have been reasonably expected, that the South African
generic and subgeneric terms are applicable to the Central African
types, hence the nine new genera described in the present paper.
Generally speaking, the family, as represented in Central
Africa, has not, so far, shown in any marked degree, the peculiar
local phases common to both genera and species such as the
author found when dealing with the agnathous forms from the
same localities, and which were described last year in these
‘ Proceedings.’
AFRICARION CONCAVOSPIRA, sp. n. (Pl. IL. fig. 20.)
Shell subovate, very thin, almost membranaceous, transparent,
polished, straw-colour; whorls 3, the first two small, sunken,
sculptured with punctate spiral striz, the last whorl large, raised
above the first two, dilated in front, marked only with transverse
growth-plications ; suture deeply impressed, bordered below by an
ill-defined granular zone; columella margin descending in a
younded curve; labrum simple, projecting in front, receding
above and below ; aperture broadly dilated.
Alt. 7:25, diam. maj. 12, diam. min. 8°5 mm.
Aperture : alt. 7°25, diam. 7°5 mm.
Hab. Kiduha, Lake Mutanda, S.W. Uganda. (Robin Kemp.)
* For explanation of the Plates see p. 810,
788 MR. H. B. PRESTON ON NEW GENERA AND
AFRICARION CoPIosA, sp.n. (PI. II. fig. 13.)
Shell subelliptical, solidly corneous, brownish olive; whorls 3,
rapidly increasing, the Jast very large, marked with transverse,
radiate growth-plice and irregular, distant, scvatch-like, spiral
sulcations; suture impressed ; base of shell moderately convex ;
columella margin arched above, angled and obliquely descending
below; labrum simple, considerably projecting in front, receding
above and below ; aperture ovate.
Alt. 9°75, diam. maj. 17, diam. min. 12°5 mm.
Aperture: alt. 9, diam. 10°5 mm.
Hab. Burunga, Mount Mikeno, Belgian Congo, at an altitude
of 6000 feet. (Robin Kemp.)
AFRICARION KIDUHAENSIS, sp. n. (PI. II, fig. 16.)
Shell differing from 4. copiosa Preston in its more globular
form, membranaceous texture, slightly paler colour, more obliquely
descending columella margin and more circular aperture.
Alt. 9, diam. maj. 14°75, diam. min. 11 mm,
Aperture: alt. 8°25, diam. 9 mm.
Hab. Kiduha, Lake Mutanda, 8.W. Uganda. (Robin Kemp.)
AFRICARION MARSABITENSIS, sp. n. (PI. II. fig. 14.)
Shell ovate, much inflated, thin, pale reddish brown ; whorls 3,
the last two rapidly increasing, the earlier whorls marked with
punctate spiral striz, the last with arcuate transverse growth-
lines only; suture very lightly impressed, narrowly margined
below; base of shell inflated; columella margin obliquely
descending, broadly curving below; labrum simple, acute ;
aperture roundly ovate.
Alt. 12°75, diam. maj. 15°75, diam. min. 12 mm.
Aperture: alt. 10°75, diam. 9°5 mm.
Hab. Northern slopes of Mount Marsabit, British East Africa,
at an altitude of 4600 feet. (A. Blayney Percival.)
AFRICARION ORESTIAS, sp. n. (Pl. II. fig. 15.)
Shell rather small, ovate, very thin, almost membranaceous,
pale yellowish brown, polished, shining; whorls 3, rapidly
increasing, the last very large, moderately convex, projecting in
front; suture almost linear, narrowly margined below; columella
margin descending in a rounded curve; aperture wide, very
large, dilated, roundly ovate; labrum membranaceous, receding
above and below.
Alt. 6, diam. maj. 9°5, diam. min. 7:25 (nearly) mm.
Aperture: alt. 5°5, diam. 5°75 mm.
Hab. Slopes of Mount Kenia, British East Africa. (Robin
Kemp.)
AFRICARION OSCITANS, sp.n. (PI. II. fig. 4.)
Shell thin, pale amber-colour, semiovate, depressed, the spire
very slightly exserted ; whorls 3, rapidly increasing, the first two
SPECIES OF ZONITIDH FROM EQUATORIAL AFRICA, 789
sculptured with spiral punctate lines, the last obsoletely, obliquely,
arcuately, transversely plicate; suture impressed, margined above,
the margin being of a pinkish colour; columella margin almost
vertically descending above, gently curved below ; labrum simple,
projecting in front, receding considerably below and a. little
above ; aperture large, wide, subovate.
Alt. 9°5, diam. maj. 13°5, diam. min. 9°75 mm,
Aperture: alt. 7, diam. 8 mm.
Hab. Lake Mutanda, 8.W. Uganda. (Robin Kemp.)
AFRICARION SPATIOSA, sp. n. (PI. I. fig. 17.)
Shell oblong, depressed, thin, nearly membranaceous, straw-
colour, polished, shining; whorls 3, the first two very small and
sculptured with somewhat distant, punctate, spiral strie, the last
very large,and marked only with radiate, wrinkle-like plications ;
suture lightly impressed, callously margined below ; base of shell
inflated ; columella excavated ; aperture ovate, very wide, and
broad.
Alt. 7°5, diam. maj. 11°75, diam. min. 8 mm.
Aperture: alt. 7°25, diam. 9°25 mm.
Hab. Burunga, Mount Mikeno, Belgian Congo, at an altitude
of 6000 feet. (Robin Kemp.)
AFRICARION KAGAMBAHENSTS, sp. n. (PI. IT. fig. 18.)
Shell differing from 4. spatiosa Preston in its more ovate form,
in the greater elevation of the spire, in its broader and much
more convex base, and in the shape of the aperture, which is much
narrower and higher in proportion to its breadth; moreover, the
punctate spirals on the earlier whorls are much finer and more
closely set than is the case with A. spatiosa.
Alt. 8 (nearly), diam. maj. 11, diam. min. 8 mm.
Aperture: alt. 7°75, diam. 8°25 mm.
Hab. Kagambah, 8.W. Uganda. (Robin Kemp.)
AFRICARION TENEPROSA, sp.n. (PI. II. fig. 19.)
Shell subelliptical, thin, almost membranaceous, dark reddish
brown, polished, shining; whorls 3, rapidly increasing, the last
large, sculptured with transverse growth-lines ; suture impressed,
margined below, the margin being of a darker colour than the
remainder of the shell ; columella margin descending obliquely,
obtusely angled and excavated below ; Tabr um simple, projecting
in front, receding above and below; aperture ovate.
Alt. 7, diam. maj. 10°25, diam. min. 9°5 mm.
Aperture: alt. 6, diam. 8 mm.
Hab. Burunga, Mount Mikeno, Belgian Congo, at an altitude
of 6000 feet. (Robin Kemp.)
GUDEELLA ARANEA, Ssp.n. (PI. IIT. fig. 20.)
Shell somewhat depressedly turbinate, yellowish brown ;
whorls 5, regularly increasing, the last subangular just above the
790 MR. H. B. PRESTON ON NEW GENERA AND
periphery; sculptured with extremely fine, closely-set, wavy,
microscopic, spiral strie, which appear to be rather coarser in the
subsutural region ; suture impressed, narrowly margined below ;
perforation narrow, internally slightly angled and sculptured with
somewhat distant and coarse, spiral lire crossed by radiate growth-
puckerings ; columella descending in a curve, very slightly out-
wardly retlexed above; labrum simple; aperture obliquely and
compressedly sublunate.
Alt. 3°25, diam. maj. 6, diam. min. 5:25 mm.
Aperture: alt. 2, diam. 2-5 mm.
Hab. Between Entebbe and Masaka, 8.W. Uganda. (Robin
Kemp.)
The internal sculpture of the umbilicus somewhat resembles
the pattern of a spider’s web, hence the trivial name.
GUDEELLA BARTAENSIS, sp.n. (PI. IIL. fig. 6.)
Shell very depressedly turbinate, pale brownish yellow ;
whorls 43, regularly increasing, sculptured throughout with very
fine, closely-set, wavy, spiral striz ; suture impressed, very nar-
rowly margined ; umbilicus narrow, deep; columella outwardly
veflexed above, very slightiy obliquely descending; labrum
simple; aperture sublunate.
Alt. 3°5 (nearly), diam. maj. 6°25, diam. min. 5:25 mm.
Aperture: alt. 3, diam. 2:75 mm.
Hab. Barta Steppes, British East Africa, (A, Blayney
Percival.)
GUDEELLA KAMPALAENSIS, sp. n. (PI. IIL. fig. 10.)
Shell differing from G. bartaénsis Preston in its smaller size,
much more narrowly margined suture, and in being microscopically
spirally striate throughout; the aperture is also more obliquely
and compressedly sublunate.
Alt. 3, diam, maj. 5°25, diam. min. 5 (nearly) mm.
Aperture: alt. 2°25, diam. 2°25 mm.
Hab. Kampala on Lake Victoria Nyanza, 8.W. Uganda; also
found at Entebbe and Jinga in the same district. (Robin Kemp.)
GUDEELLA URGUESSENSIS, sp. n. (PI. ITI. fig. 17.)
Shell closely allied to G. bartuénsis Preston, but rather broader,
with wider umbilicus and much more oblique columella; the
aperture is considerably broader, though no higher, and the
microscopic spiral sculpture is much finer than is the case in that
species.
Alt. 3°75, diam. maj. 6°75, diam. min. 6 mm.
Aperture : alt. 2°25, diam. 2:75 mm.
Hab. Urguess, British East Africa, (A. Blayney Percival.)
GUDEELLA CONSUETA, sp. n. (Pl. III. fig. 18.)
Shell thin, depressedly turbinate, polished, shining, pale reddish
brown ; whorls 44, regularly increasing, the last subangulate at
SPECIES OF ZONITIDE FROM EQUATORIAL AFRICA. 791
the periphery, sculptured throughout with very fine, wavy, spiral
strie ; suture impressed, margined below; umbilicus narrow,
deep; columella margin obliquely descending, extending above
into a thin, parietal callus; labrum acute; aperture rather
obliquely sublunate.
Alt. 3, diam. maj. 6, diam. min. 5°25 mm.
Aperture: alt. 2°5, diam. 2°5 mm.
Hab. Between Mbarara and Masaka, S.W. Uganda. (Robin
Kemp.)
GUDEBLLA usITATA, sp.n. (PI. III. fig. 2.)
Shell differing from G. consweta Preston in its smaller size and
comparatively rather more elevated spire, in having half a whorl
less, and in its more vertical columella and proportionately broader
aperture; the spiral sculpture is also much finer on the whorls
than on the base of the shell, while in G. consweta it is regular
throughout.
Alt. 2°75, diam. maj. 4°5, diam. min. 4 mm.
Aperture: alt. 2, diam. 2°5 mm.
Hab. Between Masaka and Entebbe, 8.W. Uganda. (Robin
Kemp.)
GUDEELLA DENSESCULPTA, sp. n. (PI. III. fig. 14.)
Shell very depressedly turbinate, thin, subhyaline, pale greyish
straw-colour; whorls 5, regularly increasing, very closely, micro-
scopically, spirally striate ; suture impressed, callously margined
below, the marginal callus being rather more coarsely and dis-
tantly, spirally striate than the remainder of the shell; umbilicus
narrow; columella descending in a very oblique curve ; labruin
simple; aperture rather obliquely sublunate.
Alt. 3°5, diam. maj. 7, diam. min. 6 mm.
Aperture: alt. 3, diam. 2°75 mm.
Hab. Kagambah, 8.W. Uganda. (Robin Kemp.)
GuDEELLA MIME, sp. n. (PI. IIT. fig. 23.)
Shell differing from G. densesculpta Preston in its much
smaller size, in having one whorl less, and in its darker colour,
which is of an amber hue; the last whorl is subangulate above,
the columella margin descends vertically and the aperture, though
sublunate, is less oblique.
Alt. 2. diam. maj. 3°75, diam. min. 3°25 mm.
Aperture: alt. 1°75, diam. 1°75 mm.
Hab. Kagambah, 8.W. Uganda. (Robin Kemp.)
GUDEBLLA ELGONENSIS, sp.n. (PI. III. fig. 19.)
Shell turbinate, not very depressed, polished, shining, pale
brownish horn-colour; whorls 5, regularly increasing, the last
rounded, the remainder flattish, sculptured with very fine and
closely-set, microscopic, wavy, spiral stria, the same sculpture
also occurring on the hase of the shell; suture impressed, very
792 MR. H. B. PRESTON ON NEW GENERA AND
narrowly margined below ; umbilicus narrow, partly overhung by
the outward expansion of the upper portion of the columella ;
columella margin outwardly expanded and slightly curved above,
not outwardly expanded and very obliquely descending below ;
labrum simple; aperture broadly, obliquely sublunate.
Alt. 3°75, diam. maj. 6°5, diam. min. 6 mm.
Aperture: alt, 2°75, diam. 3 (nearly) mm.
Hab. Mount Elgon, Uganda. (C. W. Woodhouse.)
GUDELLA GERSTENBRANDTI, sp. n. (PI. ITI. fig. 7.)
Shell differing from G. elgonensis Preston in its smaller size
and more compressed last whorl, darker colour, and more curved
columella ; the aperture also is proportionately not so broad and
is less oblique.
Alt. 3°5 (nearly), diam. maj. 5°75, diam. min. 5 mm.
Aperture: alt. 2°5, diam. 2°5 mm.
Hab. Mount Elgon, Uganda. (C. W. Woodhouse.)
GUDEELLA WooDHouSsEI, sp. n. (PI. III. fig. 13.)
Shell allied to G. elgonensis Preston, but larger, with more
obtuse apex and more convex whorls, the columella is much more
vertically descending and the aperture is proportionately higher
than in that species.
Alf. 5, diam. maj. 7°25, diam. min, 6°5 mm.
Aperture: alt. 3, diam. 3 mm.
Hab. Mount Elgon, Uganda. (C. W. Woodhouse.)
GUDEHLLA INCLINANS, sp. n. (PI. III. figs. 1 & 8.)
Shell depressed with sloping sides, thin, reddish brown becoming
paler on the base towards the umbilical area; whorls 4, rather
rapidly increasing, having a somewhat drooping appearance, the
last very slightly descending i in front, marked with radiate growth-
plicee which are more noticeable in the subsutural region and
sculptured throughout with fine, wavy, spiral strie ; suture im-
pressed, margined below ; umbilicus narrow, partly concealed by
the outward expansion of the columella; columella margin out-
wardly expanded and descending in a gentle curve above, oblique
below ; labrum simple, acute, projecting in front, slightly receding
above, greatly receding below; aperture broadly and somewhat
obliquely sublunate.
Alt. 3°75 (nearly), diam. maj. 7, diam. min. 6 mm.
Aperture: alt. 3°25, diam. 3 mm.
Hab. Burunga, Mount Mikeno, Belgian Congo, at an altitude
of 6000 feet. (Robin Kemp.)
GUDEELLA INFLATA, sp. n. (PI. IIT. fig. 21.)
Shell differing from G. inclinans Preston in its larger size,
much more inflated form, and darker colour which is uniform
throughout, much finer spiral sculpture and more narrow um-
bilicus, more vertically descending columella and ovate aperture,
SPECIES OF ZONITIDE FROM EQUATORIAL AFRICA, 793
Alt. 5°25, diam. maj. 7°75, diam. min. 6°5 mm.
Aperture: alt. 4:25, diam. 3°75 mm.
Hab. Barunga, Mount Mikeno, Belgian Congo, at an altitude
of 6000 feet. (Robin Kemp.)
GUDEHLLA IRIDESCENS, sp. n. (PI. III. fig. 15.)
Shell allied to @. inclinans Preston, but rather larger, though
having the same number of whorls and with coarser spiral
Sculpture, the sutural margin is broader and the columella
descends vertically above, and is then angled and very oblique
below, the aperture consequently being broader than in G. in-
clinans; when placed under the microscope the surface of the
shell appears to be highly iridescent.
Alt. 4:25, diam. maj. 7°75, diam. min. 6°20 mm,
Aperture: alt. 3°25, diam. 3°5 mm.
Hab. Mount Mikeno, Belgian Congo. (Robin Kemp.)
GUDEELLA MARSABITENSIS, sp. n. (PI. III. fig. 22.)
Shell somewhat depressedly orbicular, thin, semi-transparent,
pale reddish brown ; whorls 5, regularly increasing, the last sub-
angulate above, sculptured with very fine, wavy, closely-set, spiral
striz ; suture impressed, margined below, the margin rather more
coarsely spirally striate than the remainder of the surface and
having the spiral striz crossed by very oblique, scratch-like lines ;
base of shell moderately convex, sculptured with exceedingly fine,
microscopic, wavy, spiral stri# and coarser, somewhat distant,
revolving lines; umbilicus rather wide, deep; columella margin
outwardly expanded above, curved, obliquely descending below ;
labrum simple ; aperture obliquely sublunate.
Alt. 3°5, diam. maj. 6°75, diam. min. 6 mm.
Aperture: alt. 2°75, diam. 3 mm.
Hab. Northern slopes of Mount Marsabit, British East Africa,
at an altitude of 4600 feet. (A. Blayney Percival.)
GUDEELLA MASAKAENSIS, sp.n. (Pl. III. fig. 16.)
Shell very depressedly turbinate, olivaceous brown, polished,
shining; whorls 4, regularly increasing, sculptured with very
fine, wavy, closely-set, microscopic, spiral striz ; suture impressed,
margined below, the sculpture on the margins, as also on the base
of the shell, being slightly coarser than on the remainder of the
upper surface; umbilicus narrow, not deep; columella margin
outwardly reflexed above, descending in a very oblique curve,
extending into a very thin, but well-defined, parietal callus which
enters the mouth of the shell just behind the upper margin of
the labrum ; labrum simple, receding below ; aperture obliquely,
broadly, and rather compressedly sublunate.
Alt, 2°5, diam. maj. 5, diam. min. 4°25 mm.
Aperture: alt. 1:75, diam. 2 mm.
Hab. Between Entebbe and Masaka, S.W. Uganda. (Robin
Kemp.)
794 MR. H. B. PRESTON ON NEW GENERA AND
GUDEBLLA MULTISTRIATA, sp. n. (Pl. III. fig. 3.)
Shell small, thin, depressed, suborbicular, pale yellowish horn-
colour; whorls 4, sculptured throughout with fine, very closely-
set, spiral striae, the last large ; suture lightly impressed, margined
below ; perforation nearly concealed by the rather broad outward
expansion of the columella margin; columella margin obliquely
descending; labrum simple, acute; aperture broadly obliquely
sublunate.
Alt. 2°75, diam. maj. 5°75, diam. min. 5 mm.
Aperture: alt. 2°5, diam. 2°75 mm.
Hab. Buhamba, near Lake Kivu, Belgian Congo, (Robin
Kemp.)
Allied to Thapsia insimulans Smith * from the Nyika Plateau,
British Central Africa, but larger, with broader whorls and
coarser spiral sculpture.
JUDEELLA MUKANDANSIS, sp. n. (PI. ITI. fig. 11.)
Shell differing from G. kigeziensis Preston T in its larger size,
much paler colour, more exserted spire, narrower umbilicus, more
oblique aperture, and in being microscopically spirally sculptured
above, which is not the case with G. kigeziensis.
Alt. 4:5, diam. maj. 9°25, diam. min. 8 mm.
Aperture: alt. 4, diam. 3°75 mm.
Hab. Mukanda, near Lake Kivu. (Robin Kemp.)
GUDEELLA MUKANDAENSIS, Var. MUTANDANA, var.n. (PI. IIL.
fig. 12.)
Shell differing from the typical form in being rather more
widely perforate, darker in colour, with oblique columella margin
and more rounded aperture.
Alt. 4°75, diam. maj. 9°5 (nearly), diam. min. 7:75 mm.
Aperture: alt. 4:25, diam. 4°25 mm.
Hab. Lake Mutanda, 8.W. Uganda. (Robin Kemp.)
GUDEELLA CONSOBRINA, sp. n. (PI. ITT. fig. 9.)
Shell differmg from G. mukandaénsis Preston in its colour,
which is of a creamy hue, much more inflated form, more verti-
cally descending columella margin, less oblique, wider, and more
compressed aperture, and in the total absence of spiral sculpture.
Alt. 5, diam. maj. 9°25, diam. min. 8 (nearly) mm,
Aperture: alt. 4°25, diam. 4:25 mm.
Hab. Mukanda, near Lake Kivu. (Robin Kemp.)
GUDEELLA NEMORUM, sp. n. (PI. IIT. fig. 24.)
Shell perforate, depressedly turbinate, thin, somewhat polished,
very light reddish brown; whorls 5, regularly increasing, not
convex, the last slightly shouldered above, sculptured with very
Proc. Zool. Soc. London, 1899, p. 583, pl. xxxiii. figs. 16, 17, 18.
Proc. Malac. Soc. London, x. p. 285, 1913.
SPECIES OF ZONITIDE FROM EQUATORIAL AFRICA, 795
fine, closely-set, wavy, spiral strive ; suture impressed, margined
below ; base of shell sculptured as above ; umbilicus rather wide,
deep ; columella margin outwardly expanded, descending in a
very oblique curve, diffused above into a thin, well-defined, out-
wardly projecting, parietal callus which reaches the upper margin
of the labrum; labrum acute, slightly sinuous, projecting in
front, receding below and slightly above ; aperture rather broadly
obliquely sublunate.
Alt. 3°75, diam. maj. 7, diam, min. 6 mm.
Aperture: alt. 3, diam. 3°25 mm.
Hab. Forests to the north of Mount Kenia, British East
Africa, (A. Blayney Percival.)
GUDEELLA PALLIDIOR, sp.n. (Pl. III. fig. 5.)
Shell depressedly orbicular, greyish yellow, polished, shining ;
whorls 4, regularly increasing, sculptured with microscopic,
closely-set, spiral striz ; base of shell sculptured with fine, slightly
distant and wavy, revolving, scratch-like striz; suture impressed,
margined below; umbilicus narrow, very slightly overhung by
the outward expansion of the columella; columella margin
descending in an oblique curve; labrum simple, whitish at the
extreme edge; aperture rather broadly, obliquely, and compressedly
sublunate.
Alt. 3, diam. maj. 5°75, diam. min. 5 mm.
Aperture: alt. 2°5, diam. 2°75 (nearly) mm.
Hab. Urguess, British East Africa, (A. Blayney Percival.)
GUDEELLA TRIBULATIONIS, sp. n. (Pl. IIT. fig. 4.)
Shell depressedly turbinate, shining, pale reddish brown;
whorls 4, sculptured throughout with fine, closely-set, wavy,
spiral striz ; base of shell pale yellowish grey ; suture impressed,
narrowly margined below, the margin being of a whitish colour;
umbilicus deep, but very narrow ; columella vertically descending
above, slightly oblique, and curved below; labrum simple;
aperture compressedly sublunate. — .
Alt. 3°5, diam. maj. 5°75, diam. min. 5 mm.
Aperture: alts 2°5, diam. 2°75 (nearly) mm.
Hab. Mount Elgon, 8.W. Uganda. (C. W. Woodhouse.)
ELGONELLA, gen. nov.
Shell rather small, corneous, perforate, turbinate or depressedly
so, marked only with transverse plications or growth-strie, and
having the labrum slightly reflexed.
Genotype: LZ. euloteformis Preston.
The genus should also inelude :—
g
Zingis gaziensis Preston, from Gazi, British East Africa, Ann.
Mag. Nat. Hist. London, ser. 8, vol. vii. 1911, p. 467, pl. xi. fig. 10.
Froc. Zoou, Soc.—1914, No. LIV. 54
796 MR. H. B. PRESTON ON NEW GENERA AND
Zingis consanguinea Preston, from the Mount Kenia Region,
Ua omteses We
Zingis kempi Preston, from 8.W. Uganda, Proc. Zool. Soc.
London, 1912, p. 185, pl. xxxii. fig. 14.
ELGONELLA EULOTEFORMIS, sp.n. (Pl. II. figs. 9a, 9b, 9.)
Shell perforate, moderately solid, turbinate, pale yellowish,
slightly polished ; whorls 5, regularly increasing, sculptured with
very oblique, transverse growth plications and strie#; suture
impressed ; umbilicus somewhat wide, deep ; columella outwardly
reflexed above, descending in a rather oblique curve; labrum
acute, narrowly reflexed; aperture compressedly and very
broadly sublunate.
Alt. 5°5, diam. maj. 7°25, diam. min. 6°5 mm.
Aperture: alt. 4°25, diam. 3°25 mm.
Hab. Kagambah, 8.W. Uganda. (Robin Kemp.)
ELGONELLA BRUNNEA, sp. n. (PI. II. figs. 11a, 11 6.)
Shell perforate, turbinate, dark grey, covered with a thin, dark
brown periostracum; whorls 5, the last somewhat convex,
marked with oblique, rather coarse growth-ridges; suture im-
pressed, erenellated by the terminations of the transverse erowth-
ridges; umbilicus rather wide, deep; columella descending
obliquely, curved above and slightly outwardly expanded ; labrum
simple ; aperture broadly sublunate.
Alt. 4:75 (nearly), diam, maj. 7 (nearly), diam. min. 6 mm.
Aperture: alt. 3, diam. 2°5 (nearly) mm.
Hab. Rumruti, British East Africa. (Robin Kemp.)
ELGONELLA DISCOLORATA, sp.n. (PI. II. figs. 7a, 7 6.)
Shell depressedly turbinate, greyish, mottled, clouded and
closely spirally striated or banded with cream-colour; whorls 5,
regularly increasing, smooth but for growth-markings ; suture
impressed ; umbilicus moderately wide, deep ; columella vertically
descending above, curved below, slightly reflexed; labrum
simple ; aperture somewhat oblique, ovate.
Alt. 4:5, diam. maj. 8, diam. min. 7 mm.
Aperture: alt. 3°25, diam. 3 mm.
Hab. Larogi Hills, British East Africa. (A. Blayney Percival.)
The cream -coloured markings seem to be caused by local
opaqueness in the texture of the shell.
ELGONELLA FLAVIDULA, sp. n. (PI. II. figs. 5a, 5 6.)
Sheli perforate, depressedly turbinate, rather thin, pale yel-
lowish ; whorls 5, marked only with irregular, oblique, arcuate
growth-lines ; suture impressed ; umbilicus rather broad, open
deep; columella margin outwardly expanded above, descending
in a curve; labrum simple; aperture compressedly ovate.
Alt. 5, diam. maj. 8°25, diam. min. 7:25 mm.
Aperture ; alt. 3°25, diam. 2°75 mm.
Hab. Mount Elgon, Uganda. (C. W. Woodhouse.)
SPECIES OF ZONITIDE FROM EQUATORIAL AFRICA, 797
ELGONELLA ANGUSTIOR, sp. n. (PI. IT. figs. 6a, 66.)
Shell allied to H. favidula Preston, but of a pale brown colour
and more narrowly turbinate, with subangulate last whorl; the
umbilicus is narrower, the aperture is higher and narrower, and
the margins of the labrum are convergent.
Alt, 4°5, diam. maj. 7°25, diam. min. 7 mm,
Aperture: alt, 3, diam. 3 mm,
Hab. Mount Elgon, 8.W. Uganda. (C. W. Woodhouse.)
ELGONELLA onIBATES, sp. n. (PI. II. figs 8a, 8b.)
Shell small, turbinate, light reddish brown, smooth, slightly
polished ; whorls 43, regularly increasing, marked with oblique
growth-lines; suture impressed; umbilicus very narrow, deep;
columella rather broadly outwardly expanded, descending in a
rather gentle curve ; labrum simple; aperture roundly ovate.
Alt. 4°25, diam. maj. 5:75, diam. min. 5 mm.
Aperture: alt. 3, diam. 3 mm.
Hab. Mount Elgon, Uganda. (C. W. Woodhouse.)
ELGONELLA ROBINI, sp.n. (Pl. I. figs. 10a, 10 6.)
Shell perforate, turbinate, light brown, smooth, but for oblique
growth-ridges ; whorls 43, regularly increasing, the last subangu-
Jate at the periphery; suture impressed; umbilicus moderately
narrow, deep; columella slightly excavated above, somewhat
vertically descending below; labrum simple; aperture very
compressedly, broadly sublunate.
Alt. 4, diam. maj. 6°25, diam. min. 5°75 mm.
Aperture ; alt. 2°75, diam. 2°5 mm.
Hab. Rumruti, British East Africa. (Robin Kemp.)
ELGONELLA SOBRINA, Sp.n. (PI. IT. figs. 12a, 126.)
Shell differing from #. robini Preston in its smaller and pro-
portionately much more depressed form and in its rounded last
whorl; it has half a whorl less and the growth-ridges are much
more marked and the umbilicus is narrower; the columella and
the aperture are somewhat more oblique and the latter is pro-
portionately broader.
Alt. 3°5, diam. maj. 5°75, diam. min. 5 mm.
Aperture: alt. 2°25, diam. 2°5 mm.
Hab. Rumruti, British Kast Africa. (Robin Kemp.)
BURUNGAELLA, gen. nov.
Shell thin, corneous, whorls rather tightly coiled, the last
proportionately large, more or less obsoletely, transversely plicate ;
aperture somewhat large.
Genotype: &. oscitans Preston.
BURUNGAELLA OSCITANS, sp. n. (PI. I. figs. 17a, 176, 17.)
Shell perforate, very thin, almost membranaceous, very de-
pressedly turbinate above, convex below, pale brownish straw-
798 MR. H. B. PRESTON ON NEW GENERA AND
colour; whorls 5, regularly increasing, the last large, marked
with obsolete, oblique, transverse, crease-like plications which are
more developed in the subsutural region ; suture impressed ; base
of shell somewhat inflated; umbilicus rather narrow, deep,
partly overhung by the outward expansion of the columella;
columella very slightly curved, scarcely oblique; labrum simple,
membranaceous, projecting a little in front ; aperture ovate.
Alt. 7:5, diam. maj. 10°75, diam. min. 8°25 mm.
Aperture: alt. 6°25, diam. 4°5 mm.
Hab. Burunga, Mount Mikeno, Belgian Congo, at an altitude
of 6000 feet. (Robin Kemp.)
BuRUNGAELLA BUHAMBAENSIS, sp. n. (PI. I. fig. 16.)
Shell perforate, thin, corneous, turbinate, pale yellowish brown;
whorls 5, the first small, the second somewhat large in proportion,
the remainder regularly increasing, slightly convex, irregularly
and obsoletely plicate and sculptured with wavy, transverse strie,
the last whorl angled at the periphery; suture impressed, very
narrowly margined below ; umbilicus very narrow, deep, almost
covered by the upper, broad, outward expansion of the columella
margin; columella margin membranaceous, broadly, triangularly,
outwardly expanded and nearly vertically descending above,
then angled and very obliquely descending below; aperture
broadly and rather compressedly obliquely subcrescentie.
Alt. 5°5, diam. maj. 7, diam. min. 6 mm.
Aperture: alt. 4, diam. 3 mm.
Hab. Buhamba, near Lake Kivu, Belgian Congo. (Robin
Kemp.)
BURUNGAELLA IMPERFORATA, Sp. n. (PI. I. fig..15.)
Shell imperforate, turbinate, very thin, yellowish brown; whorls
5, regularly increasing, the last somewhat inflated, the last two
microscopically spirally striate and marked with oblique, trans-
verse, crease-like plicee ; suture impressed ; base of shell inflated ;
columella obliquely descending, outwardly, callously expanded
above; aperture depressedly and very broadly sublunate.
Alt. 6°25, diam. maj. 9°25, diam. min. 7°75 mm.
Aperture: alt. 4°75, diam. 4°25 mm.
Hab. Burunga, Mount Mikeno, Belgian Congo, at an altitude
of 6000 feet. (Robin Kemp.)
BURUNGAELLA MUTANDANA, sp.n. (PI. I. fig. 14.)
Shell rather depressedly turbinate, thin, pale yellowish brown;
whorls 43, regularly increasing, the last subangulate at the
periphery, marked with oblique transverse riblets or creases, which
become obsolete on the base; suture impressed, crenellated by
the terminations of the transverse riblets; umbilicus moderately
narrow, deep; columella rather broadly outwardly expanded
SPECIES OF ZONITIDA FROM: EQUATORIAL AFRICA, 799
above, descending in a somewhat cblique curve; labrum thin,
almost membranaceous, very slightly reflexed; aperture ovate.
Alt. 5, diam. maj. 8°75, diam. min. 7°25 mm,
Aperture; alt. 4, diam. 4°25 (nearly) mm.
Hab. Lake Mutanda, extreme 8.W. Uganda. (Robin Kemp.)
BLAYNEYELLA, gen. nov.
Shell thin, corneous, narrowly umbilicate, depressedly turbinate,
transversely plicate; labrum a little reflexed.
Genotype: &. percivali Preston.
Generally, but not always, ornamented with a peripheral
purplish band.
Zingis papyracea Preston, from 8.W. Uganda, Proc. Zool. Soe.
London, 1912, p. 185, pl. xxxii. fig. 15, may also be considered as
belonging to Blayneyella.
BLAYNEYELLA PERCIVALI, sp. n. (Pl. I. figs. 19@, 196, 19.)
Shell perforate, turbinate, thin, pale yellowish brown, painted
with a rather narrow, supersutural, reddish-purple band, which
appears on the last whorl as a superperipheral band; whorls 5,
the last gradually descending in front, marked with oblique,
arcuate, irregular, crease-like riblets; suture impressed, faintly
margined below; umbilicus moderately narrow, very deep, slightly
overhung by the outward reflexion of the columella; columella
eurved, diaphanous, rather broadly outwardly reflexed above,
narrowly so below; labrum simple; aperture ovate; interior of
shell showing the superperipheral band through the test.
Alt. 9°5, diam. maj. 14°25, diam. min. 11°75 mm.
Aperture: alt. 7°25, diam. 6-75 mm.
Hab. Larogi Hills, British East Africa, (A. Blayney Percival.)
BLAYNEYELLA KISENGIENSIS, sp.n. (PI. I. fig. 21.)
Shell perforate, depressedly turbinate, thin, very pale reddish
brown; whorls 54, regularly, but rather rapidly, increasing,
marked with somewhat indistinct, oblique, radiate riblets; suture
impressed; base of shell rather convex; umbilicus moderately
wide, deep; coiumella outwardly expanded above, descending in
a curve; aperture broadly sublunate.
Alt. 8°25, diam. maj. 13°25, diam. min. 11 mm,
Aperture: alt. 5°25, diam. 5°75 mm.
Hab. Kisengi, Lake Kivu, German East Africa, at an altitude
of 4500 feet. (Robin Kemp.)
BLAYNEYELLA PURPURHOCINGTA, sp.n. (PI. I. fig. 20.)
Shell perforate, thin, corneous, depressedly turbinate, yellowish
brown, painted with a narrow, supersutural, reddish-purple band,
which appears on the last whorl as a superperipheral band ;
whorls 6, regularly increasing, marked with oblique transverse
800 MR. H. B. PRESTON ON NEW GENERA AND
growth-strie ; suture impressed; base of shell somewhat inflated ;
umbilicus moderately wide and deep; columella rather outwardly
expanded, descending in a curve, oblique below; labrum thin,
slightly reflexed ; aperture very broadly sublunate.
Alt. 6°5, diam. maj. 11:25, diam. min. 9°25 mm.
Aperture: alt. 5°5, diam. 5 mm.
Hab. Nairobi, British East Africa. (A. Blayney Percival.)
BLAYNEYELLA MICROSPIRALIS, sp. n. (PI. I. fig. 18.)
Shell allied to B. purpureocincta Preston, but differing from
that species in its much more depressed form, more obliquely
descending columella, and narrower and more compressed aperture,
and finely spirally sculptured first whorl, which sculpture is only
visible by the aid of the microscope.
Alt. 6, diam. maj. 10°25, diam. min. 8-5 mm.
Aperture: alt. 4°5, diam. 4°5 mm.
Hab. British East Africa.
LAROGIELLA, gen. nov.
Shell umbilieate, thin, corneous, turbinate, transversely plicate ;
labrum membranaceous.
Genotype: Z. venatoris Preston.
To the above genus should also be assigned :—
Natalina permembranacea Preston, from 8.W. Uganda, Proc.
Zool. Soc, London, 1912, p. 183, pl. xxxi. figs. 20, 20 a, & 205.
Zingis planispira Preston, also from S.W. Uganda, t.c. p. 185,
pl. xxxil. fig. 16.
LAROGIELLA VENATORIS, sp.n. (PI. I. figs. 22, 226, 22.)
Shell somewhat depressedly turbinate, thin, yellowish brown ;
whorls 57, regularly increasing, the last angled at the periphery
and descending in front, marked with oblique growth-plicee ;
suture impressed, almost imperceptibly margined below ; umbilicus
moderately wide, deep, slightly overhung by the triangular, out-
ward expansion of the columella margin; columella margin
descending in a curve; labrum membranaceous; aperture com-
pressedly ovate.
Alt. 7-5, diam. maj. 12°5, diam. min. 10-25 mm.
Aperture: alt. 5, diam. 5-5 mm.
Hab. Larogi Hills, British East Africa, at an altitude of from
6000 to 7000 feet. (A. Blayney Percival.)
LAROGIELLA ANGULIFERA, sp. n. (PI. II. fig. 1.)
Shell perforate, depressedly conic, thin, pale brown, painted
with an indistinct, whitish, infrasutural band; whorls 5, the
last descending in front, regularly increasing and angled at the
periphery, marked with irregular, oblique, arcuate, transverse,
crease-like riblets and showing traces of spiral whitish lines,
SPECIES OF ZONITIDA FROM EQUATORIAL AFRICA, 801
especially in the subsutural region, which latter appear to be in
the texture of the shell and not at all in the nature of outward
sculpture ; suture impressed ; base of shell slightly convex, rather
more noticeably marked with the whitish textural lines than the
rest of the shell; umbilicus moderately wide, deep; columella
diffused upwards into a very thin, well-defined, and sinuous
parietal callus, which reaches the upper margin of the labrum,
curved and outwardly reflexed above, oblique and scarcely reflexed
below; labrum simple; aperture oblique, compressedly subovate,
Alt. 8°75, diam. maj. 13°75, diam. min. 11-25 mm.
Aperture: alt. 5°5, diam. 5°75 mm.
Hab. Larogi Hills, British Kast Africa. (A. Blayney Percival.)
LAROGIELLA FONTICULA, sp.n. (PI. II. fig. 3.)
Shell perforate, thin, roundly turbinate, fulvous; whorls 5,
regularly increasing, sculptur ed with irregular, very oblique,
radiate, transverse plications and microscopically shagreened ;
suture impressed ; umbilicus narrow, deep, well-like, a little over-
hung by the outward expansion of the columella margin ;
columella margin vitreous, broadly outwardly expanded above,
descending in a curve ; 3 labrum simple ; aperture roundly ovate.
Alt. 6°75, diam. maj. 10, diam. min. 8 mm.
Aperture: alt. 5, diam. 4-5 mm.
Hab. Lake Mutanda, 8S.W. Uganda. (Robin Kemp.)
LAROGIELLA KOMBAENSIS, sp.n. (PI. II. fig. 2.)
Shell perforate, thin, submembranaceous, pale yellowish brown ;
whorls 4, regularly increasing, sculptured throughout with regular,
oblique, somewhat arcuate, transverse, crease-like plice; suture
impressed; base of shell moderately inflated ; umbilicus rather
wide, deep; columella vitreous, widely outwardly reflexed above,
descending in a very slightly oblique curve; labrum thin, acute,
projecting a little in front; aperture ovate.
Alt. 4°75, diam. maj. 10, diam. min. 8 mm.
Aperture: alt. 5°25, diam. 4 mm.
Hab. Komba, 8.W. Uganda. (Robin Kemp.)
LAROGIELLA MALASANJIENSIS, sp. n. (PI. I. fig. 23.)
Shell somewhat depressedly turbinate, thin, pale brown; whorls
5, regularly increasing, the last subangulate at the periphery,
marked with oblique, arcuate, transverse plications; suture im-
pressed; base of shell marked with very fine, wavy, revolving
striz in.addition to the transverse sculpture ; umbilicus moderately
wide, deep; columella descending in an oblique curve, outwardly
expanded above; labrum thin, membranaceous; aperture broadly
and compressedly sublunate.
Alt. 5°25, diam. maj. 9°5, diam. min. 8°25 mm.
Aperture: alt. 4°5, diam. 3°25 mm.
Hab, Malasanji, 8.W. Uganda. (Robin Kemp.)
802 MR. H. B. PRESTON ON NEW GENERA AND
NAKURUELLA, gen. nov.
Shell umbilicate, turbinate, thin, corneous, transversely plicate,
with broad aperture.
Genotype: Zingis bullata Preston *.
NaAkURUELLA soror, sp.n. (PI. I. figs. 13a, 136, 13¢.)
Shell differing from WV. bullata Preston in its more depressed
form, in having : a whorl less, in its still narrower umbilicus, and
sisiilive more oblique and much broader a perture; the basal
portion of the labrum is also slightly reflexed.
Alt. 11, diam. maj. 17, diam. min. 13 mm.
Aperture: alt. 8°5, diam. 8°75 mm.
Hab. Burunga, Mount Mikeno, Belgian Congo, at an altitude
of 6000 feet. (Robin Kemp.)
MIKENOBLLA, gen. nov.
Shell thin, corneous, narrowly perforate, with somewhat elevated
spire, transversely plicate only, labrum acute.
Genotype: J. ahena Preston.
Species which should probably be referred to this genus are :—
Zingis gregorii Smith, from Mt. Kenia, Proc. Malac. Soe.
Lond. vol. i. p. 164.
Zingis aurea Preston, from British East Africa, Rev. Zool.
Africaine, Bruxelles, vol. iii. p. 47, pl. v. fig. 6.
MIKENOELLA AHENA, sp. n. (PI. II. figs. 26 a, 265, 26.)
Shell perforate, conically turbinate, very thin, pale yellowish
brown; whorls 5, regularly increasing, the last large, obsoletely
transversely plicate, the plicee being more noticeable in the sub-
sutural region ; suture impressed, very narrowly margined below ;
base of shell inflated ; utubilicus narrow, deep, partly concealed
by the outward reflexion of the columella margin; columella
margin broadly and membranaceously outwardly reflexed above,
almost vertically descending in a gentle curve; labrum acute ;
aperture ovate.
Alt. 11-25, diam. maj. 13, diamn. min. 11 mm.
Aperture: alt. 7°25, diam. 5 mm.
Hab. Burunga, Mount Mikeno, Belgian Congo, at an altitude of
6000 feet. (Robin Kemp.)
MIKENOSLLA ELEVATA, sp.n. (PI. IJ. fig. 25.)
Shell perforate, inflatedly turbinate, thin, yellowish brown,
painted just above the suture and periphery with a broad spiral
band of purplish red, below which occurs a peripheral narrow
band of a creamy colour; whorls 53, regularly increasing, the last
inflated, marked only, especially on the upper whorls, with fine
oblique growth-plice; suture impressed; base of shell inflated,
# Rey. Zool. Africaine, Bruxelles, yol. ui. pp. 47-48, pl. v. fig. 4.
SPECIES OF ZONITIDA: FROM EQUATORIAL AFRICA, 803
marked with radiate arcuate growth-plications and discoloured
with a purplish tinge in the circum-umbilical region ; perforation
narrow, deep, slightly overhung by the outward expansion of the
columella; columella triangularly outwardly expanded above,
descending in a gentle curve, the margins slightly converging,
receding below; aperture subovate.
Alt. 11, diam. maj. 12°5, diam. min. 10°25 mm.
Aperture: alt. 6, diam. 5-75 mm.
Hab. Uarogi Hills, British Hast Africa, at an altitude of
between 6000 and 7000 feet. (A. Blayney Percival.)
MIKENOBLLA NEGLECTA, sp. n. (PI. II. fig. 27.)
Shell globosely turbinate, narrowly perforate, thin, unicolorous,
pale yellowish brown; whorls 53, regularly increasing, marked
only with slightly arcuate, transverse growth-plice ; base of shell
convex; umbilicus very narrow, almost covered by the outward
reflexion of the columella; columella broadly outwardly reflexed
above, very narrowly so below; aperture subovate.
Alt. 11, diam. maj. 13°25 (nearly) mm.
Aperture: alt. 7-5, diam. 6°5 mm.
Hab. Kiduha, $.W. Uganda. (Robin Kemp.)
URGUESSELLA, gen. nov.
Shell perforate, thin, corneous, depressed, hirsute, the last
whorl descending rather rapidly in front.
Genotype: U. urguessensis Preston.
The following two species should undoubtedly be included in
Urguessella :—
Trachycystis nigrotincta Preston, from British Hast Africa, Rev.
Zool. Africaine, vol. i. p. 325, pl. xvii. fig. 9.
T. fusco-olivacea Smith, from Nyassaland, Proc. Zool. Soe.
London, 1899, p. 585.
URGUESSELLA URGUESSENSIS, sp. n. (PI. III. figs. 25, 25 a, 25 6.)
Shell perforate, turbinate, with subplanulate spire, yellowish
brown, covered with a hirsute periostracum, the hairs being about
one millimetre in length, yellow in colour, placed at a moderate
distance apart and arranged in very oblique rows; whorls 43,
regularly increasing, the last descending in front; suture 1m-
pressed ; base of shell somewhat convex ; umbilicus rather narrow,
deep; columella triangularly outwardly expanded above, de-
scending in a rounded curve; labrum white, thin, very narrowly
outwardly expanded and reflexed; aperture somewhat rect-
angularly ovate, on the parietal wall; just without the aperture,
where the hairs no longer appear, the surface of the shell is seen
to be marked with transverse pustulous riblets.
Alt. 5:25, diam. maj. 9°25, diam. min. 8 mm.
Aperture: alt. 3°75, diam. 4 mm.
Hab. Urguess, British East Africa. (A. Blayney Percival.)
804 MR. H. B, PRESTON ON NEW GENERA AND
URGUESSELLA ESAU, sp. n. (PI. III. fig. 26.)
Shell allied to U. urguessensis Preston, but differing con-
siderably from that species in its thinner texture, smaller size,
more reddish colour, less planulate spire, subangulate last whorl,
and much finer and shorter hairs, which are arranged in very
oblique rows, but running at a much less acute angle to the
suture than in U. wrguessensis; the columella is also higher and
more angularly curved, while the aperture is roundly ovate.
Alt. 4°75, diam. maj. 9, diam. min. 7‘25 mm.
Aperture: alt. 3°75, diam. 3°75 mm.
Hab. Urguess, British East Africa. (A. Blayney Percival.)
URGUESSELLA CAPILLATA, sp. n. (PI. III. fig. 28.)
Shell suborbicular, depressed, with almost planulate spire, thin,
pale reddish brown; whorls 4, regularly increasing, the last
descending in front, sculptured with corrugated, closely-set,
obliquely arcuate, transv erse plice, bearing at intervals broad
bristly hairs; suture impressed; base of shell slightly convex ;
umbilicus moderately wide, deep; columella broadly outwardly
reflexed above and diffused into a thin, ill-defined, glassy, parietal
callus, descending in a vertical curve; aperture obliquely sub-
lunate.
Alt. 3°25, diam. maj. 7, diam. min. 6 mim.
Aperture: alt. 3, diam. 2°5 mm.
Hab. Burunga, Mount Mikeno, Belgian Congo, at an altitude
of 6000 feet. (Robin Kemp.)
URGUESSELLA CUTICULARIS, sp. n. (PI. ITI. fig. 27.)
Shell thin, suborbicular, with depressed spire, covered with a
shortly hispid, dark Neowen periostracum ; whorls 43, regularly
increasing, marked with very oblique, arcuate, transverse plica-
tions, the last whorl angulate at the periphery ; suture impressed ;
base of shell rather inflated ; umbilicus rather broad, open, deep ;
columella broadly outwardly expanded above, descending in an
oblique curve ; labrum simple; aperture subrectangular.
Alt. 4, diam. maj. 7, diam. min. 6 mm.
Aperture: alt. 3°5, diam. 3 mm.
Hab. Malasanji, 8.W. Uganda. (Robin Kemp.)
TROCHOZONITES BUHAMBAENSIS, sp. n. (Pl. II. fig. 23.)
Shell acutely conic, pale brown, somewhat polished ; whorls 7,
slightly convex, the first three somewhat slowly increasing in
breadth, smooth, the remainder rather rapidly increasing, sculp-
tured with coarse, oblique, transverse riblets, the last whorl
strongly carinate at the periphery ; suture impressed, very
narrowly margined; base of shell nearly smooth; umbilicus
narrow and very deep, partly concealed by the reflexion of the
columella margin ; columella margin outwardly expanded above,
SPECIES OF ZONITIDA FROM EQUATORIAL AFRICA. 805
vertically descending ; labrum acute, simple; aperture irregularly
subrectangular.
Alt. 7, diam. maj. 6°25, diam. min. 5°25 mm,
Aperture: alt. 2°75, diam. 2°25 mm.
Hab. Buhamba, near Lake Kivu, Belgian Congo. (Robin
Kemp.)
TROCHOZONITES EXPATRIATA, sp. n. (PI. II. fig. 21.)
Shell conical, with obtuse apex and very slightly constrieted
sides, very thin, reddish brown, glossy ; whorls 6, the last two
rather convex, the last acutely carinate at the periphery, sculp-
tured with course, oblique growth-ridges ; suture faintly incised,
narrowly eallously margined above ; base of shell nearly planulate ;
umbilicus very narrow, almost entirely concealed by the outward
expansion of the columella; columella margin descending in a
very slight and very oblique curve; labrum sunple; aperture
subrectangular.
Alt. 6, diam. maj. 6°5, diam. min. 6 mm.
Aperture: alt. 1-5, diam. 3 mm.
Hab. Burunga, Mount Mikeno, Belgian Congo, at an altitude
of 6000 feet. (Robin Kemp.)
In form much resembling the species of the genus inhabiting
West Africa.
TROCHOZONITES KEMPI, sp. nu. (PI. II. fig. 22.)
Shell moderately small, conic, with very laterally compressed
spire, in dead condition yellowish grey, somewhat iridescent,
polished, shining; whorls 64, the first three and a half convex,
the remainder flattened, sloping, the last acutely carinate at the
periphery, sculptured with closely-set, oblique, transverse strive ;
suture linear, very narrowly margined above; base of shell
slightly convex; umbilicus narrow, deep; columella somewhat
outwardly expanded, obliquely descending; labrum simple ;
aperture curvedly subrectangular.
Alt. 5:25, diam. maj. 6°75, diam. min. 6°5 mm.
Aperture: alt. 2, diam. 3 mm.
Hab. Buhamba, near Lake Kivu, Belgian Congo. (Robin
Kemp.)
TROCHOZONITES SUTURALIS, sp.n. (PI. II. fig. 24.)
Shell rimate, turritely turbinate, somewhat laterally compressed
below, thin, olivaceous; whorls 61, regularly increasing, smooth
but for very oblique, transverse growth-lines ; suture impressed,
bearing, especially on the lower part of the shell, an erect, soine-
what callously thickened ridge, which appears as a carina on the
last whorl; perforation very narrow, almost concealed by the
exceedingly narrow outward reflexion of the columella ; columella
thickened, outwardly reflexed and vertically descending above,
obliquely descending below, very slightly inwardly bulging in the
806 MR. H. B. PRESTON ON NEW GENERA AND
median region ; labrum acute, receding below, advancing above ;
aperture subrectangular.
Alt. 7, diam. maj. 6°25, diam. min. 6 mm.
Aperture: alt, 3, diam. 2°5 mm.
Hab. Urguess, British East Africa. (A. Blayney Percival.)
PERCIVALIA, gen. nov.
Shell thin, perforate, compactly coiled, turbinate, sculptured
on the spire with transverse wrinkle-like plications, which are
absent on the bases.
Genotype: P. nyiroénsis Preston.
PERCIVALIA NYIROENSIS, sp.n. (Pl. L. figs. 12 a, 12d.)
Shell depressedly turbinate, polished, shining, pale greenish
yellow, painted with a broad, supersutural, spiral band of chestnut,
which appears as a superperipheral band on the last whorl ;
whorls 6, the first two slightly exserted, the remainder flattish,
the last angular at the periphery, sculptured with oblique, trans-
verse, arcuate, wrinkle like plice; suture impressed, narrowly
margined above; base of shell somewhat convex, sculptured only
with very fine, indistinct, wavy, spiral striz; umbilicus very
narrow, partly concealed by the twisted columella; columella
somewhat sharply twisted above, very obliquely descending ;
labrum simple; aperture rather obliquely sublunate.
Alt. 6, diam. maj. 10, diam. min. 9°25 mm.
Aperture: alt. 5, diam. 4°75 mm.
Hab. Mount Nyiro, to the south of Lake Rudolph, at an
altitude of 8300 feet. (A. Blayney Percival.)
LEDOULXIA CRASSIPLICATA, sp.n. (PI. I. figs. 1, 1a.)
Shell depressedly turbinate, thin, light brown; whorls 5,
regularly increasing, convex, shouldered above, sculptured with
coarse, rather distant, obliquely arcuate, somewhat acute plications,
the last whorl acutely and callously carinate at the periphery ;
suture impressed, narrowly callously margined above; base of
shell moderately convex, smooth, without sculpture ; umbilicus
open, deep, sculptured with somewhat distant, wavy, revolving
strie within ; columella margin broadly outwardly expanded and
vertically descending above, curved below; labrum simple ;
aperture obliquely subrectangular.
Alt. 7, diam. maj. 11:5, diam. min. 10 mm.
Aperture: alt. 5, diam. 4°5 mm.
Hab. Forest to the north of Mount Kenia, British East Africa.
(A. Blayney Percival.)
LEDOULXIA DECUSSATA, sp. n. (PI. I. fig. 7.)
Shell with roundly turbinate spire, pale brownish yellow above,
the colouring concentrating in an ill-defined, supersutural, reddish-
brown band, showing traces of having been covered with a
SPECIES OF ZONITIDE FROM EQUATORIAL AFRICA, 807
chestnut periostracum; whorls 6, regularly increasing, the
extreme apex smooth, the second whorl sculptured only with
slightly distant and very oblique, coarse, transverse stiie, the
remaining whorls sculptured with fine, closely-set, transverse
striz crossed by fine spirals, thus presenting a finely decussate
appearance, the last whorl rather strongly carinate at the
periphery ; base of shell polished, shining, dark brownish yellow,
somewhat granulate and sculptured with fine, wavy, moderately
closely-set, revolving strie; suture impressed, very narrowly
callously margined above; umbilicus rather narrow, deep;
columella outwardly expanded in the upper region, vertically
descending above, then bulging obliquely inwards and very gently
and obliquely curved below; labrum simple; aperture broadly
and somewhat compressedly sublunate.
Alt. 11, diam. maj. 16, diam. min. 14°75 mm.
Aperture: alt. 7, diam. 7°75 mm.
Hab. Larogi Hills, British East Africa. (A. Blayney Percival.)
LEDOULXIA ELGONENSIS, sp.n. (PI. I. fig. 4.)
Shell allied to Martensia jenynsi Pfr.*, of which it may
ultimately prove to be a subspecies; it is, however, much more
depressed than is that species, the spiral sculpture, both on the
base and on the whorls, is also more developed, and the columella
descends much more obliquely.
Alt. 7°75, diam. maj. 13, diam. min. 11:25 mm.
Aperture: alt. 5°75, diam. 5°75 mm.
Hab. Mount Elgon, Uganda. (C. W. Woodhouse.)
LEDOULXIA EUSSOENSIS, sp.n. (PI. I. fig. 2.)
Shell rather solid, perforate, somewhat depressedly turbinate,
cream-coloured, flecked here and there with greyish black and
ornamented with a very narrow, interrupted, peripheral band of
the same; whorls 6, sculptured with very closely-set, coarse,
oblique, arcuate striz; the last whorl obtusely angled at the peri-
phery ; base of shell marked with lines of growth and sculptured
with irregular, wavy, revolving, scratch-like striz ; suture well
impressed ; umbilicus narrow, “deep, slightly overhung by the
outward expansion of the Golumella: columella descending in a
curve; labrum simple; aperture obliquely sublunate.
Alt. 13°75, diam. maj. 19, diam. min. 16°5 mm.
Aperture: alt. 9, diam. 8 mm.
Hab. Eusso Nyiro, British East Africa. (Robin Kemp.)
LEDOULXIA JINGAENSIS, sp.n. (Pl. I. fig. 6.)
Shell broadly turbinate, thin, straw-colour, painted with a
narrow, supersutural, spiral band of reddish purple ; whorls 6,
regularly i increasing, rather convex, the last acutely and callously
carinate at the periphery ; sculptured with fine, closely-set,
* Proc. Zool. Soc. London, 1845, p. 131.
808 ME. H. B. PRESTON ON NEW GENERA AND
oblique, slightly arcuate, transverse costule ; suture impressed,
narrowly callously margined above; base of shell moderately
convex, somewhat polished, marked with lines of growth and
sculptared with fine, wavy, revolving strie; umbilicus narrow,
deep; columella descending in an oblique curve, outwardly ex-
panded above and diffused into a very thin, almost imperceptible,
well-defined, parietal callus, which reaches the upper margin of
the labrum; labrum simple, acute; aperture obliquely, com-
pressedly sublunate.
Alt, 9, diam. maj. 14, diam. min. 12°25 mm.
Aperture: alt. 5°75, diam. 6°25 mm.
Hab. Jinga on Lake Victoria Nyanza, 5.W. Uganda; also
taken at Entebbe and Kampala in the same district. (Robin
Kemp.)
LEDOULXIA LEVISTRIATA (Preston*), var. NYERIENSIS, var. 0.
(Bi ne: 3.)
Shell differing from the typical form in its rather broader
shape and proportionately slightly less acuminate spire, and in
the colour, which in the present variety is of a yellowish hue.
Alt. 12, diam. maj. 19°75, diam, min. 17 mm.
Aperture: alt. 8°5, diam. 9°5 mm.
Hab. Nyeri, British East Africa, (Robin Kemp.)
LEDOULXIA ADJACENS, sp.n. (PI. I. fig. 8.)
Shell differing from Z. levistriata (Preston) in its more depressed
form, rather narrower perforation, and slightly broader aperture ;
the transverse sculpture on the whorls is finer and the spiral
sculpture on the base rather coarser and considerably more
distant ; the colour of the present species is yellowish brown, and
the carina and sutural margination not so marked as in JZ, levi-
striata.
Alt. 11°5, diam, maj. 18, diam. min. 15°5 mm.
Aperture: alt. 8, diam. 8°25 mm.
Hab. Mount Kenangop, Aberdare Range, British East Africa.
(Robin Kemp.)
LEDOULXIA MARSABITENSIS, sp. n. (PI. I. fig. 5.)
Shell allied to Martensia permanens Smithy, but smaller,
darker in colour, more narrowly umbilicate, more closely and
coarsely sculptured, and with proportionately higher spire than
has that species.
Alt. 9°75, diam. maj. 15, diam. min. 13°5 mm.
Aperture: alt. 7, diam. 6°25 mm.
Hab. Northern slopes of Mount Marsabit, British East Africa.
(A. Blayney Percival.)
x * Martensia levistriata Preston, Rev. Zool. Africaine, Bruxelles, iii. p. 48, pl. v.
g. 2.
“+ J. Malac. viii. p. 94.
SPECIES OF ZONITIDE FROM EQUATORIAL AFRICA, 809
FALLOONELLA, gen. nov.
Shell depressed or depressedly conoid, perforate, closely and
almost laminiferously striate above, polished on the base, suturally
callously margined, and carinate at the periphery.
Genotype: /. exquisita Preston.
FALLOONELLA EXQUISITA, sp. n. (PI. I. figs. 9, 9 a, 96.)
Shell depressedly turbinate, suborbicular, thin, somewhat
diaphanous and shining, chestnut-coloured ; sino 5 1, the earlier
whorls smooth, the later whorls sculptured with moderately fine
and very closely-set, oblique, arcuate costule; suture bearing a
sulcate raised ridge, which appears on the last whorl asa yellowish
raised peripheral carina; base of shell polished, shining, showing
radiate growth-ridges and sculptured only with very fine, wavy,
and closely-set revolving striz; umbilical area greenish olive,
the perforation being deep and very narrow ; columella oblique,
somewhat curved, very narrowly outwardly expanded above, and
diffusing into a thin, minutely granular, outwardly spreading,
well-defined callus, which reaches the upper margin of the labrum ;
labrum simple, acute ; ; aperture obliquely and somewhat br oadly
sublunate.
Alt. 5, diam. maj. 9°75, diam. min. 8°75 mm,
Aperture: alt. 4:25, diam. 4°5 mm.
Hab. Urguess, British Kast Africa. (A. Blayney Percival.)
FALLOONELLA EXQUISITA GUDEI, subsp. n. (PI. I. fig. 10.)
Shell differing from typical /. exquisita Preston in having one
whorl more, in its less depressed form, rather finer cblique
costule on the spire, and in the sutural ridge which is also con-
siderably finer, the perforation is rather less narrow, the columella
is less curved and more oblique, and the aperture is rather
narrower in proportion to its height.
Alt. 6°75, diam. maj. 11 (nearly), diam. min. 9°75 mm.
Aperture: alt. 5°25, diam. 5 mm.
Hab. Larogi Hills, British Hast Africa. (A. Blayney Percival.)
FALLOONELLA LAROGIENSIS, sp.n. (PI. I. figs. lla, 116.)
Shell perforate, depressedly turbinate, dark livid flesh-colour
above, covered with a chestnut periostracum ; whorls 54, flattish,
regularly, but not rapidly, increasing, the ‘first two and a half
spirally sulcate, the remainder sculptured with moderately coarse,
not very resular, but wavy, oblique, transverse riblets, presenting
through a lens a slightly corrugated appearance ; suture lightly
impressed, bearing a somewhat coarse, thread-like, cream-coloured
ridge, which appears as a rather sharp peripheral carina on the
last whorl; base of shell slightly polished, whitish yellow, sculp-
tured with distant, revolving, microscopic, scratch-like striz ;
erforation narrow, deep; columella triangularly, outwardly ex-
panded above, obliquely descending in a slight curve; labrum
810 MR. H. B. PRESTON ON NEW GENERA AND
simple, receding below ; aperture obliquely and depressedly
crescentic,
Alt. 6, diam. maj. 11°75, diam. min. 10 mm.
Aperture : alt. 5:25, diame 5 mm.
Hab. Larogi Hills, British East Africa, at an altitude of from
6000 to 7000. feet. (A. Blayney Percival.)
EXPLANATION OF THE PLATES.
Pram Te
Hess eem solace Ledoulwia crassiplicata, sp. n.
2 + eussoensis, Sp. 1.
3. “- levistriata (Preston), var. nyeriensis, var. 1.
4, + elganensis, Sp. 0.
5 3 marsabitensis, Sp. i.
6 +) Jingaénsis, sp. 1.
is bs decussata, sp. 0.
8. 7 adjacens, sp. Nn.
9,9a,9b. Falloonella exquisita, sp. n.
10. a A gudei, subsp. n.
iil ays Wallop 4 lavogiensis, sp. N.
124,126. Percivalia nyiroénsis, sp. n.
13.4,136,13¢. Nakuruélla soror, sp. n.
14, Burungaélla mutandana, sp. 0.
15. or imperforata, sp. 0.
16. 3% buhambaénsis, sp. 0.
17 a, 176, 17 ¢. Bs oscitans, sp. N.
18. Blayneyella microspiralis, sp. n.
19 a,19 6,19. Ff percivali, sp. n.
20 5 purpureocincta, sp. N.
21. xy kisengiensis, Sp. ll.
22,22 6, 22c. Larogiella venatoris, sp. n.
23. malasanjiensis, Sp. i.
Prate II.
Figs il Larogiella angulifera, sp. n.
2. a kombaénsis, sp. ii.
3. ” fonticula, sp. n.
4, Africarion oscitans, sp. N.
5a,5b. EHlgonella flavidula, sp. n.
6a, 6b. a angustior, sp. 1.
Wl i, Wh (i 5 discolorata, sp. n.
8a, 8b. 2 oribates, sp. 0.
9a,9b, 9c. 50 euloteformis, sp. ni.
10a, 106. 9 robini, Sp. 0.
lla, 116. “3 brunned, sp. N.
12a, 12.6. sobrina, sp. 0.
13. Afr icarion copiosa, sp. 0.
14, 3 marsabitensis, sp. N.
15. 5 orestias, sp. Nn.
16. 6 kiduhaénsis, sp. .
ie o spatiosa, sp. n.
18. = kagambahensis, sp. n.
19. x tenebrosa, sp. n.
20. concavospira, Sp. I.
21. Trochozonites expatriata, sp. n.
22. 53 kempi, sp. 0.
23: at buhambaeénsis, sp. 0.
24. suturalis, sp. n.
25. Mikenoélla elevata, sp. Nl.
26 a, 26 b, 26.¢. . ahena, sp. 0.
27. BA neglecta, sp. nl.
St ee.
SPECIES OF ZONITIDA FROM EQUATORIAL AFRICA,
5.
Figs.
2 VS OU go bo
a
[2
i
$2 NS? OU Go bo
to bo bo bo
SSS
24,
Prate III,
Gudeélla inclinans, sp. n.
33
usitata, sp. n.
multistriata, sp. 1.
tribulationis, sp. n.
pallidior, sp. n.
bartaénsis, sp. n.
gerstenbrandti, sp. n.
inclinans, sp. n.
consobrina, sp. ni.
kampalaénsis, sp. n.
mukandaénsis, sp. i.
var. mutandana, var. n.
woodhousei, sp. n.
densesculpta, sp. n.
iridescens, sp. Nn.
masakaénsis, sp. n.
urguessensis, Sp. N.
consueta, Sp. N.
elgonensis, sp. n.
araned, sp. n.
inflata, sp.n.
marsabitensis, sp. n.
mime, sp. Nn.
nemorum, sp. n.
25, 25 a, 25 5. Urguessella urguessensis, Sp. ni.
26.
27,
28.
esau, Sp. n.
cuticularis, sp. n.
capillata, sp. n.
Proc. Zoou. Soc.—1914, No. LV. 55
811
wn
P.Z.S.1914. BOULENGER. 2k.
J. Green photo del.
1. ATELOPUS SPURRELLI. 2. LEPIDOBLEPHARIS INTERMEDIUS.
3. POLYCHRUS SPURRELLI.
at
abt be t
P.Z.S.1914 . BOULENGER, Bile
J. Green photo del.
2, HOMALOCRANIUM NIGRUM.
1 LEPTOPHIS BREVIOR:.
3h JEVL/MIES) SIP ORR VEIL IC I
ON BATRACHIANS AND REPTILES FROM COLOMBIA,
813
44. On a second Collection of Batrachians and Reptiles
made by Dr. H: G. F. Spurrell, F.Z.S., in the Choco,
Colombia. By G. A. BouLencer, F.RS., F.Z.8.*
[Received May 20, 1914; Read June 9, 1914. ]
(Plates I. & II.+)
INDEX.
Page
Geographical Zoology: Choco, Colombia, Batrachians and
Systematic :—
Atelopus spurrelli, sp. n.
Lepidoblepharis intermedius, sp. NM. ......... see ceeeee eee cee ees
JEQUGCHUFOIS SLU RAND S05 Wa ses 420 decaas S50ce0 Bac do0kod 630800 50cod
JEG DRO DOIS: WRBTTOR, Sido TWasscace o6ho00 e50n60 06800600 ee dad agaGon cae
JOT MHDORTRIOUD TOOGIPOHOs SDS 25 coc doo Lou nooiscco09 050090090 GooHo
J HIGDS CP DOUIRGUUG:, S061 book née cca dendsedon sou occ eee see op seEo Hepa dTORC
813
813
814
814
815
816
817
Since the publication of the account of Dr. Spurrell’s Collection
of Batrachians and Reptiles from the Choco, the British
Museum has received a further important series of these animals
presented by the same energetic collector. All were obtained at
or near Pena Lisa, Condoto. In the following list I have only
enumerated the species not mentioned in the previous list.
BATRACHIA.
CAUDATA.
1. SPELERPES PARVIPES Peters.
ECAUDATA.
2. BuFo CONIFERUS Cope.
3. HyYLopEs RANIFORMIS Bler.
4, ATELOPUS SPURRELLI, sp. n. (PI. I. fig. 1.)
Habit slender. Head a little longer than broad, one-third the
len
eth from snout to vent; snout obtusely pointed, prominent,
obliquely truncate, a little longer than the eye; loreal region
nearly vertical, slightly concave; nostril near the tip of the
snout ; interorbital space as broad as the upper eyelid.
limb slender, as long as the trunk; fingers webbed at the base,
with swollen tips; first finger very short; a flat palmar tubercle ;
no subarticular tubercles, Hind limb slender; the tibio-tarsal
* Published by permission of the Trustees of the British Museum.
+ For explanation of the Plates see p. 817.
+ P.Z.S. 1913, p. 1019.
)
5*
Fore
814 MR. G. A. BOULENGER ON BATRACHIANS
articulation reaches the eye; tibia half the length of head and
body ; toes half-webbed, inner very short but perfectly distinct ;
the tips blunt; metatarsal and subarticular tubercles very
indistinct. Green above, with small black spots and large
symmetrical black markings, viz.: a streak on the canthus
rostralis; an X on the head and between the shoulders, the
anterior branches extending on the upper eyelids, the posterior
confluent with a lateral band which is expanded on the temple
and extends to the groin; a spade-shaped figure on the sacral
region; a pair of large spots above the waist, and cross-bars on
the limbs; lower parts white, with round black spots. Male with
brown rugosities on the inner finger.
From snout to vent 24 mm.
A single specimen.
REPTILIA.
CHELONIA.
1. CHELYDRA ROSSIGNONI Bocourt.
This species was only known from Southern Mexico, Guatemala,
and Western Ecuador.
LACERTILIA.
2. THECADACTYLUS RAPICAUDA Houtt.
3. LEPIDOBLEPHARIS INTERMEDIUS, sp. n. (PI. I. fig. 2.)
Scaling as in LZ. peracce Blgr., form more as in L. feste Peracea.
Granules of upper parts very small, largest on the snout. Snout
pointed, much longer than the orbit; rostral with median cleft,
and concave above, as in the two other species; symphysial
likewise very large, with two clefts behind; five upper and four
lower labials, first very large. Brown above, with darker and
lighter variegations; a whitish streak across the nape ; lower
parts pale brown, throat whitish.
Motaleleng thie. on...) seen, eeeeees 63 mm.
ISIGAVUL Sons coc eee ee annem 8" 5,
Wradithvotaheads. 4... 45, ei Ge ee
TBACGIWfe\ Seen, cree ee ee ee 2 Joes
ore shim lyases. eats San;
Liman tees Batraoys. ates. a Tel
TOTES 2 Co Skee es ere meaner 34,
Two specimens.
4. ANOLIS PALMERI Bley.
5. PoLYCHRUS SPURRELLI, sp.n, (PI. I. figs. 3, 3a.)
Snout obtusely pointed ; nostril equally distant from the orbit
and from the tip of the snout ; eye-opening nearly as large as the
AND REPTILES FROM COLOMBIA. 815
tympanum : upper head-scales smooth or feebly striated ; scales of
supraorbital semicircles in contact with each other in the middle
or separated by one series of scales; supraocular scales small ;
labials striated, five upper and four lower to below centre of eye;
symphysial with a median cleft posteriorly. Gular scales much
larger than ventrals, feebly striated: no gular denticulation or
erest. Scales on body feebly unicarinate, dorsals a little larger
than laterals and ventrals. Limbs moderately elongate, the
scales feebly unicarinate. 12 to 15 femoral pores on each side.
Tail very long and round, with rather strongly unicarinate scales.
Reddish or purplish brown, posterior part of belly and anal region
greenish ; A-shaped darker cross-bands may be present on the
body ; a blackish horizontal line behind the eye and another,
oblique, from below the centre of the eye; base of tail and base of
thigh with a white streak.
Mo taller et ligesaenersn teem a i: 360 mm.
SE ICA\G aaocee Seas cach neee agen ere nao D3) |)
Width of head ......... SR SOR eae.
IB Od, Sea ceeeaecesien eu h yatta palsies One
Roney limnily irae aun at verre Nar. 5s ON res
IB Ghee lian) Oeea so ie One Aneel cts Seas
ARs ees ney ease aerate aoe ean QO
Two female specimens.
6. IGUANA TUBERCULATA Laur.
OPHIDIA.
7. Boa mmeERATOR Daud.
8. EPIcRATES CENCHRIS L,
9. DRYMOBIUS BODDAERTII Sentz.
10. Purynonax Pa@crLonotus Gthr.
11. Spinores PuLLATUS L.
12. SprmorEs MEGALOLEPIS Gthr.
This rare snake is only known from N.W. Ecuador and
S.W. Colombia, where specimens were obtained by Mr. Palmer.
13. Cotuser corais L.
14. LepropHis BREvioR, sp.n. (PI. II, figs. 1, 1a.)
Rostral a little broader than deep, just visible from above ;
internasals as long as broad, a little shorter than the preefrontals ;
frontal once and three-fourths as long as broad, as long as its
distance from the end of the snout, shorter than the parietals ;
816 MR. G. A. BOULENGER ON BATRACHIANS
nasal elongate, entire ; no loreal; przefrontal in contact with the
second and third upper labials ; one preocular, in contact with
the frontal ; two postoculars; temporals 1 + 2; eight upper
labials, fourth and fifth entering the eye; five lower labials
in contact with the anterior chin-shields, which are shorter than
the posterior. Scales in 15 rows, finely striated and strongly
keeled, the two outer rows smooth. Ventrals feebly angulate
laterally, 139; anal divided; subcaudals 127. Scales partly
green partly brownish, with golden sheen; upper lip white; a
black line on each side of the head, above the labialis, passing
through the eye; lower parts pinkish, with mother-of-pearl
sheen.
Total length 580 mm.; tail 245.
A single female specimen.
Well distinguished, among the species with keeled scales and
no loreal, by the low number of ventral shields.
15. XENODON COLUBRINUS Gthr.
16. RwApInmA DECORATA Gthr.
A young specimen, referred with some doubt to this species.
17. Oxyruopus PeroLartius L,
18. RuiNoBotHRYUM LENTIGINOSUM Scop.
19. OXYBELIS BREVIROSTRIS Cope.
20. OXYBELIS ACUMINATUS Wied.
21. HoMALOcRANIUM NIGRUM, sp.n. (PI. II. figs. 2, 2 a.)
Eye two-fifths the length of the snout. Rostral nearly twice as
broad as deep, scarcely visible from above; internasals half as
long as the preefrontals ; frontal pentagonal, a little longer than
broad, more than twice as broad as the supraocular, a little
longer than its distance from the end of the snout, much shorter
than the parietals; nostril between the nasals, the posterior in
contact with the preocular ; a single postocular ; temporals 1+1 ;
seven upper labials, third and fourth entering the eye ; first lower
labial in contact with its fellow behind the symphysial; four
lower jabials in contact with the anterior chin-shields, which are
longer than the posterior. Scales in 15 rows. Ventrals 143 ;
anal divided ; subcaudals 63. Black above and beneath ; a pair
of round yellow spots close together on the occiput; a larger,
oblique spot on each side of the latter, behind the angle of the
mouth; a narrow, oblique yellow spot on the fifth and sixth
upper labials, and a very small round one on the second upper
labial,
Total length 175 mm. ; tail 48.
AND REPTILES FROM COLOMBIA, 817
22. ELAPS SPURRELLI, sp.n. (PI. IT. figs. 3, 3a.)
Hye nearly as long as its distance from the mouth. Rostral
much broader than deep, scarcely visible from above; frontal
broader than the supraocular, once and a half as long as broad,
longer than its distance from the end of the snout, shorter than
the parietals ; latter as long as their distance from the end of
the snout ; one pre-and two postoculars ; temporals 1+1; seven
upper labials, third a little deeper but not larger than the fourth,
third and fourth entering the eye ; four lower labials in contact
with the anterior chin-shields, which are as long as the posterior.
Scales in 15 rows. Ventrals 232; anal divided; subcaudals 36.
52 black annuli separated by white areas on the body and by red
on the tail; the first annulus, on the nape, much broader than
the others, which are narrower than the white interspaces on the
belly and broader on the back ; a few small black spots between
the black rings on the back; sides of head, as far back as the
second temporal and the middle of the parietal, black, with a
white spot on the second, third, and fourth upper labials ; a white
middle line on the head, forming a { with a broad white bar
across the occiput.
Total length 230 mm. ; tail 20.
A single female specimen.
23. LACHESIS SCHLEGELIL Berth.
EXPLANATION OF THE PLATES.
Prate I.
10h, il Atelopus spurrelli, p. 813. Nat. size, upper and lower views.
2. Lepidoblepharis intermedius, p. 814. 14 nat. size.
3,3a. Polychrus spurrelli, p.814, Side view of head and anterior part of body,
nat, size, and upper view of head, 13 nat. size.
Prats IT,
Figs.1, la. Leptophis brevior, p. 815. Upper view of head and anterior part of
body, nat. size, and side view of head, 2.
2,2a, Homalocranium nigrum, p. 816. Upper view of head and anterior part
of body, X 2, and side view, X 3.
3,3. Hlaps spurrelli, p.817. Upper and lower views of head and anterior
part of body, X 2, and side view, X 3.
P.Z.S. 1914, CUNNINGTON. PI.
W.A.C, photo. London Stereoscopic Co., imp.
1-3. LEBRNAEOGERA] DIGE RACEPRALA:
Acts. es HAR LOGE rinAlAe 8, 9 LilEMNOGCEFPH AEA
ON PARASITIC EUCOPEPODA FROM TANGANYIKA. 819
45. Zoological Results of the Third Tanganyika Expedition,
conducted by Dr. W. A. Cunnington, 1904-1905.—
Report on the Parasitic Hucopepoda. By Winiiam A.
Cunnineton, M.A., Ph.D., F.Z.S.
[Received May 4, 1914: Read June 9, 1914. |
(Plate I.* & Text-figure 1.)
INDEX.
Systematic : Page
Lerneocera: List of described Species with synonyms ...... 820
Derneocera: Key to Species of ............ cc ccceeeceeeeeeeeeeeeee 824
DLerneocera diceracephala, Sp. Ve... 2.0... .cc eevee eee eee eee, 824
IES GH NIOCG HLH Gy Oo Ty ooo occacacn Sooscneeocayseadoncdoanadeeascengens eY40)
JE UAT HOH Ds SDo Me cooatioanse6soracnoncsere nan qonccdnesacsesoscasen | tsYAU/
1. Introduction.
In addition to the parasitic Copepods belonging to the order
Branchiura, the collections made during the Third Tanganyika
Expedition contain a very few specimens of parasitic Hucopepoda
belonging to the family Lerneide ?. While there are many
parasitic forms of Eucopepoda, comparatively few of them have
been found on freshwater hosts, and these, perhaps, have
received less attention than the forms infesting marine fishes.
In the family Lerneide, the genus Lernewocerat is the only
one which is known to occur in fresh water, and it is to this
well-known genus that our specimens have been referred.
Through the kindness of Dr. Calman I have been allowed
to examine a good many examples of Lerneocera from the Nile,
which belong to the British Museum §, Since these specimens
have not been examined or described, and since they add con-
siderably to the scanty material which we possess from the
African continent, an account of them is included in the present
paper. So far as I am aware, the existence of the genus in
Africa has never been put on record before, the species hitherto
known being either European or American. The following is
* For explanation of the Plate, see p. 829.
+ Certain parasitic Eucopepoda belonging to the family Ergasilide were obtained
by the Expedition, in addition to the forms described in this paper. They were
taken in the last free stage, in tow-nettings associated with non-parasitic Copepods,
and in consequence were dealt with by Prof. G. O. Sars in his paper on the Copepoda
of the Third Tanganyika Expedition (Proc. Zool. Soc. 1909, p. 63).
t+ The generic name is written throughout in the form in which it is almost
universally quoted, and no¢ as it was originally spelled by Blainville, viz. Lerneocera.
The word is derived from the Linnzan genus Lernea.
§ By the courtesy of the authorities of the Berlin Museum, specimens of
L. cyprinacea and LL. esocina (the latter, one of the original examples studied by
von Nordmann) were lent to the British Museum for the purpose of comparison
with the forms described here. I am particularly indebted to Dr. EK. Vanhoften for
the trouble he has taken in the matter.
820 DR. W. A. CUNNINGTON ON PARASITIC
a list of the African forms described for the first time in this
paper :—
LAKE TANGANYIKA.
Lerneocera diceracephala.
Lerneocera haplocephala.
River NILe.
Lerneocera haplocephala.
Lerneocera temnocephala.
For the purpose of illustration, I have made use of photo-
micrographs taken from the actual specimens themselves. They
are by no means easy objects to photograph, but in spite of
imperfections, the figures will make clear the various external
characters which have been used for the systematic descriptions.
I have to thank Prof. Dendy, of King’s College, London, for -
permission to do this photographic work in his laboratory.—The
plan has been adopted of giving in each case a view of the head
and cephalic arms from above. This was done by von Nordmann,
one of the earliest writers on the genus, but his example has
not been widely copied. It needs some trouble to support the
specimens in the position necessary to secure such a view, but
the figures obtained illustrate the nature of the head region
far better than any others could do.
2. Systematic Notes and Description of New Species.
The literature which deals with the different species of the
genus Lerncocera is somewhat scattered and not always easy to
obtain. Although a list of the known species with synonyms is
given by Bassett-Smith*, it is marked by material inaccuracies,
so that it would seem worth while at this point to include a list
which may be useful to future investigators as a starting point
for their researches. No attempt has been made to give an
exhaustive list of the authors by whom the species have been
cited, as this would take up a good deal of space and serve no
useful purpose. Only those works are referred to which have a
bearing on the synonymy or which contain a record of original
observations.
List of described Species with Synonyms.
|. LerN&OcERA CYPRINACEA T (Linneus) f.
‘‘Lernea tentaculis quatuor: duobus apice lunulatis.”
Linneus, Fauna Suecica, Ed, I. 1746, p. 367, tab. 11.
Lernea cyprinacea Linneus, Systema Nature, Ed. X. 1758,
p- 695.
* Proc. Zool. Soc. 1899, p. 480.
+ So far as I can ascertain, no one has ever fixed the type species or genotype of
Lerneocera. Assuming this to be the case, in order to maintain the usage of all
modern writers, I hereby select cyprinacea as genotype of Lerneocera.
~ Parentheses enclosmg the Author-citation after specific names are used in
accordance with Art. 23 of the International Rules of Nomenclature.
EUCOPEPODA FROM TANGANYIKA, 821
Lernéocera cyprinacea Blainville, Journal de Physique, t. 95,
1822, p. 377.
Lernwocera cyprinacea Burmeister, Nova Acta Acad. Ces.-
Leop. Bd. 17, 1835, p. 309.
2. LernmocerA EsocINA * Burmeister.
Lerneeocera cyprinacea v. Nordmann, Mikrograph. Beitr.
Naturgesch. wirbellosen Thiere, Heft 2, Berlin, 1832, p. 123
(non L. cyprinacea Linn.).
Lernceocera esocina Burmeister, Nova Acta Acad. Czs.-Leop.
Bd. 17, 1835, pp. 309 & 312.
Lerncocera gasterostet Brihl, Mitt. K. K. zool. Inst. d. Uniy.
Pest, 1860 (Wien), p. 1.
Lerneocera gobina Claus, Wirzb. naturw. Zeitschr. Bd. i.
IstOlle jos Mule
Lerneocera esocina Claus, Beobachtungen wher Lerneocera,
Peniculus und Lernea. Marburg, 1868, p. 1.
3. LERNHOCERA CRUCIATA Lesueur.
Lerneocera cruciata (? Lernecenicus) Lesneur, Journ. Acad. Nat.
Sci. Philadelphia, vol. 11. 1824, p, 286.
4, LERNZOCERA PHOXINACEA Kroyer.
Lerneocera phoxinacea Kollar MS., Kroyer, Naturhistorisk
Tidsskrift, ser. 3, vol. 11., Copenhagen, 1863-64, p. 399.
5. LERNHOCERA LAGENULA Heller.
Lerneocera lagenula Heller, Reise der Novara
(Wien, 1865), p. 246.
6. LERNHOCERA POMOTIDIS Krgyer.
Lerncocera pomotidis Kroyer, Naturhistorisk Tidsskrift, ser. 3,
vol. 11., Copenhagen, 1863-64, p. 397.
Crustaceen
“I
LERNXOCERA CATOSTOMI Krgyer.
Lerneocera catostomi Kroyer, Naturhistorisk Tidsskrift, ser. 5,
vol. i1., Copenhagen, 1863-64, p. 395.
1t is perhaps well to point out here that the classification of the
parasitic Eucopepoda has hardly received the attention devoted
to that of free-living forms. At the same time it is clear that
a satisfactory basis on which to classify the former is unusually
dificult to find, on aecount of the extraordinary degree of
modification commonly undergone by the female on the adoption
of a parasitic mode of life. It is possible, indeed probable, that
* It has been suggested by some modern writers (cf. Bassett-Smith, op. cit. p. 480,
and Brian, ‘Copepodi Parassiti dei Pesci d'Italia,’ Genova, 1906, p. 79) that the species
esocina and cyprinacea should be united. ‘Their contention does not seem to rest
on personal observations, but on their interpretation of the original descriptions.
After an examination of the actual specimens, [ have no hesitation im confirming the
view of the older authors, namely that the species are perfectly distinct.
822 DR, W. A. CUNNINGTON ON PARASITIC
individual variations of form will be more than usually common
as concerns the greatly distended bodies or the cephalic processes
for attachment to the host, as the exact shape would seem
without significance for the life of the parasite. Yet it is
precisely such details which are employed for the purposes of
classification. Thus within the limits of the genus Lerneocera
itself it is difficult to be sure how far the specific characters
employed will prove constant and therefore trustworthy, for
even among the specimens that I have examined a considerable
lack of uniformity has been observed,
Genus LernocerA Blainville.
It does not seem desirable to re-define the genus here,
although the species now included in it would not strictly come
under Blainville’s original generic description. Indeed, his
account is based upon certain misconceptions, notably the view—
shared by contemporary writers—of the absence of appendages
on the body, for he says ‘‘ Aucune trace d’appendices au corps.”
Nevertheless a number of species have been placed in this
imperfectly defined genus, but if is open to question whether
they should all remain there. A careful study of these forms has
given me the impression that two or three of them may merit
separation as distinct genera, or at least sub-genera; but without
opportunities for a more comprehensive examination, it is im-
possible to express a very definite opinion, and the course least
open to objection is to leave matters as they are.
Before proceeding to give descriptions of the new species, there
remain one or two matters which need some explanation. It is
characteristic of most of the species, including those which are
described in this paper, that they exhibit the peculiar boot-like
shape of the terminal portion of the body which was first referred
to by von Nordmann* in his account of Z. esocina. This is
produced, in the first place, by a protuberance immediately in
front of the genital apertures, which forms the “ heel,” and which
we may call the pre-genital prominence. In the second place,
there is generally a dorsal curvature of the hindmost portion of
the body (posterior to the genital apertures and corresponding to
the abdomen according to Claus), which, owing to the lateral
torsion undergone by the hinder part of the body, comes to lie
on one side or other of the mid-line and represents the “‘ toe.”
This explanation of the appearance we owe to Claus‘, but the
matter is made yet clearer by the conceptions on torsion in the
Lerneide quite recently put forward by Quidort. The latter
assumes that the torsion is the direct result of the mode of
fixation of the parasite and the mechanical reaction of the
external medium. Admitting the probability of this statement,
and admitting that the orientation of a parasite to its host is
* Op. cit. p. 124.
7+ Vide “ Beobachtungen tiber Lerneocera,” etc., p. 2.
= Comptes Rendus Acad. Sci. Paris, Tome 154, 1912, p. 87.
EUCOPEPODA FROM TANGANYIKA. 823
probably constant for a given species, we are furnished with an
explanation of the otherwise perplexing fact that this lateral
torsion may be either to right or left. It will be the one or the
other according to the particular side of the host which formed
the point of attachment for the parasite. Quidor, moreover,
gives evidence for believing that the amount of torsion is constant
for a given species, and can be used as a character of systematic
value.
The appendages appear to show comparatively minor differences
within the limits of this genus, and have not been appealed to
for the purpose of establishing new species. Thus I have not
deemed it necessary to study in detail the head appendages of
my new forms, since these are by no means easy to investigate,
and my material, with one exception, was very seanty. So far
as I have been able to make out, there are no points of striking
_ difference in any of the types from the arrangement which is
usual in the group. Accordingly in the specific descriptions
which follow, no special mention of head appendages, Swimming-
feet or furcal appendages is made, it being implied that these
are present in the normal manner, without baving any bearing
on the distinctions between the species.
Tn addition to the photographs reproduced in the plate, the
accompanying text-figure is given, showing in outline for the
three new species the appearance of the head and cephalic arms
from above.
Text-figure |.
Diagram showing cephalic arms, as viewed from above.
A. Lern@ocera diceracephala. LB. L. haplocephala. C. L. temnocephala.
In order to facilitate identification, and in order to emphasise
in very concrete form the chief features which characterise the
new species, a key to all the known species of Lerneocera has
been prepared. This did not prove a very casy task, as unfortu-
nately it has been possible for me to examine specimens of only
two of the forms which have been described. For particulars
of the remaining species I have been dependent entirely upon the
descriptions and figures of the authors concerned, and in such
cases no more can be done than to repeat certain statements
which would seem of value for key-making. Thus I am not
vesponsible for the rather remarkable assertion that LZ. catostomi
possesses three cephalic arms, which, of course, renders the head
quite asymmetrical. That is a feature which enables us to
824 DR. W. A. GUNNINGTON ON PARASITIC
contrast the form sharply with the typieal species of Lerncocera,
and which might serve as a claim to more than specific dis-
tinction.
It might be well to indicate here, briefly, the other forms
which in my judgment differ materially from the more normal
members of the genus. ‘The species L. lagenula, as described
and figured by Heller, retains in a great measure the primitive
segmentation of the body which is usually lost, and at the
same time fails to show the pre-genital prominence and charac-
teristic boot-like shape of the posterior end. It has also under-
gone a very slight amount of torsion. The North American
form L. pomotidis shows, according to the figure, a complete
absence of any torsion, though otherwise it might rank as_a
typical Lerneocera. Finally, it must be admitted that the form
to be described below as L. diceracephala exhibits certain features
which are non-characteristic, and one feature which is not
shared by any other species of the genus. This peculiarity, to
which reference is made in the specific name, is the existence
of only two cephalic horns—apparently the dorsal pair—ainstead
of four. Beyond this, the lobed nature of the body, suggestive of
segmentation, and the apparent absence of any torsion, are
further points of distinction.
Key to the Species of LERNZOCERA.
a. Single pair of cephalic arms present ...........0..0000ce:csseeeeesee. iceracephala.
a’. Cephalic arms markedly asymmetrical, three in number ...... catostomi.
a’. Two pairs of cephalic arms present.
b. Cephalic arms simple without any indication of forking.
c. Ventro-lateral tubercles present behind junction of
ALIS PATIO MO CaVuaR GAS eee eee ante aew select sina dectressee mentees
ce’. No ventro-lateral tubercles present.
d. Cephalic arms very long and straight; body
terminating in three broadly-rounded tubercles. pomotidis.
d’. Cephalic arms of moderate length, curved for-
wards; body terminating in five rounded
haplocephala.
_ tabercles oie eee eects eetieeterreees Oructata.
b’. Cephalic arms forked.
c. Pre-genital prominence absent ...............cccee eee Lagenula.
c’. Pre-genital prominence present.
d. Dorsal cephalic arms simple; ventral arms with
ROCESS MRR ee erence eset Pr ula eatamntat das sa sinhc om
d’. Dorsal cephalic arms forked ; ventral arms simple.
e. Ventral cephalic arms nearly as stout as
dorsal arms; egg-sacs oval, +4 length of
HOKaGl\//sapasees obs terfonn oat ek masmcebe cba cemebance Camhee
e’. Ventral cephalic arms much more slender
than dorsal arms.
Ff. Dorsal cephalic arms T-shaped ; egg-
sacs cylindrical, 3-4 length of body ... eyprinacea.
J’. Dorsal cephalic arms Y-shaped......... temnocephala.
phoxinacea.
esocina.
1. LERN#ZOCERA DICERACEPHALA, sp.n. (PI. I. figs. 1-3.)
Description.—(Adult female.) Cephalic arms only two in
number, of considerable length and projecting laterally from the
region of the head-tubercle at about right angles to the body.
EUCOPEPODA FROM TANGANYIKA. 825
The arms are dilated distally and bear, at about one-third their
length from the end, a stout postero-dorsal process which 1s
bluntly pointed. The body is bent dorsally through a consider-
able angle at a little less than one-half its length from the head.
It is not uniform in diameter, but shows a marked constriction at
about the middle and an otherwise irregular contour which may
indicate disappearing segmentation. Pre-genital prominence not
very conspicuous, simple or slightly bilobed ; terminal portion of
body not upturned and rotated very little, if at all. ‘The egg-sacs
are long and tapering, about two-fifths length of body. They
contain from four to five rows of eggs at their widest part. The
eggs are slightly oval, "14:12 mm.
Total length of complete specimen (excluding egg-sacs),
8:4 mm.*
Length of longer egg-sac, 3°5 mm.
Remarks.—The two specimens on which this new species 1s
founded are, unhappily, neither of them quite perfect. One of
them, in fact, only consists of the main part of the body, without
head, cephalic arms, or egg-sacs. This very Incomplete individual
affords, however, valuable evidence in certain respects, for the
sharp bend in the body and the noticeable constriction referred
to above, are equally recognisable here, so that we may assume
them to be definite features of the species. The specimen on
which the description mainly rests has lost a portion of one
cephalic arm, but we may fairly suppose it to have been the same
as the one which is whole. In text-fig. 1, the missing part has
been restored for the sake of affording a comparison with the
other species. It is open to question how far the lobed nature of
the body, which is so conspicuous, 1s indicative of segmentation.
The fact that the most typical members of the genus show the
body dilated into an almost formless sac, might suggest that we
are dealing in the present case with a less modified condition. On
the other hand, the position of the four pairs of swimming-feet,
which appear to have no definite relation to the body lobes, is an
argument against such a belief.
The reference in the foregoing description to the terminal
portion of the body, sufficiently indicates that in this form there
is also complete or almost complete absence of the characteristic
torsion of the body. This fact is, of course, equally displayed by
the position of the swimming-feet, which are visible in a ventral
view approximately in the middleline. All this implies, further,
that the hinder end of the body cannot exhibit the usual boot-like
shape as a consequence of deflection and rotation, as is generally
the case, and yet it must be granted that the appearance in this
respect is fairly typical. There is, of course, an important
difference, namely that the shape in question is visible only in a
lateral view, whereas it is shown in a ventral view of the more
normal types as a result of the body torsion. In the present
* This is the actual measurement of the specimen without taking into consideration
its bent state. It would measure more if straightened out.
826 DR. W. A. CUNNINGTON ON PARASITIC
instance, then, the effect is produced rather by an incision in the
region of the genital apertures than by the combination of
characters which has been already fully explained.
The most striking characteristic of this species is the existence
of only one pair of cephalic arms instead of two pairs. There
seems little doubt from the relation they bear to the head-tubercle,
that these correspond to the dorsal cephalic arms of species in
which two pairs of arms are present. Such an important differ-
ence from the common type might be considered sufficient to
warrant a generic distinction for this species, but [have preferred
to leave it for the present in the genus Lerneocera.
Occurrence.—Sumbu, Lake Tanganyika, 13.10.04. From gill-
arches of a large Clarias mossambicus. ‘Two specimens, one very
incomplete.
9. LERNHOCERA HAPLOCEPHALA, sp.n. (PI. I. figs. 4-7.)
Description.—(Adult female.) Cephalic arms four, of about
equal size, short and stout, without any indication of forking and
being so placed as to form a particularly regular cross. The
dorsal arms are simple and bluntly pointed ; the ventral differ
from them only slightly, exhibiting an obvious swelling on their
ventral aspects. The body is almost straight, unsegmented and
cylindrical. The anterior third is slender, the body dilating
gradually behind to become about twice as thick. Pre-genital
prominence well marked, simple and not bilobed ; terminal
portion of body rather slightly upturned and rotated through
somewhat less than 90°. Immediately behind the junction of the
arms with the body, and just external to the second pair of
swimming-feet, a pair of rounded tubercles are situated, which
project ventro-laterally. The egg-sacs are moderately long and
tapering, about one-fifth length of body. They contain from four
to five rows of eggs at their widest part. The eggs are approxi-
mately round, and their diameter is about *] mm.
Total length of largest specimen (excluding egg-sacs), 14°3 mm.
Remarks.—This species is represented in my material by a
considerable number of specimens from different sources, which
would suggest that it isa relatively common form. Unfortunately,
the bulk of the specimens have suffered severely from lack of care
in preservation, having been preserved apparently in the same
manner as their host, or even with it, and they are in consequence
greatly shrunken and shrivelled. Jt is thus the more satisfactory
that all these can be readily identified as belonging to this species
by the presence of the characteristic ventro-lateral tubercle
mentioned above. The single specimen from Tanganyika, on
which the description is largely based, and which is figured on
Plate I., is unluckily devoid of egg-sacs, but this defect is made
good in one from the Nile (fig. 7), which happens to be damaged
elsewhere. The Tanganyika specimen proves also considerably
loager than any of the individuals from the Nile, which is doubt-
less accounted for in part by the contracted nature of the latter.
EUCOPEPODA FROM TANGANYIKA. 827
In two tubes, each containing a number of Nile specimens, the
lengths vary from 6°2—9:0 mm. and from 8°8-11:7 mm.
A careful examination showed the lateral torsion of the body
to vary in direction, as was expected. Unfortunately, minute
details as to the position of the parasites on the host are missing
for the Nile material, so that it is impossible to test the accuracy
of Quidor’s assumptions in relation to this species. The in-
dividual from Tanganyika was, however, taken from the soft
region at the junction of the pelvic fins,—a spot more nearly
ventral than lateral. At the same time, it is hardly probable that
the point of attachment was so strictly median as to preclude the
possibility of lateral torsion in conformity with this view.
Fig. 7 serves also to show how strikingly these parasitic forms
may in turn be covered by other organisms. In this case, the
latter are Vorticellids, which infest many of these Lerneids from
the Nile to such a degree as to render difficult the study of their
anatomy. Among a considerable number of specimens taken
on a Polypterus senegalus, almost all are infested, some of
them as markedly as the one photographed. The region where
the Vorticellids are most thickly attached is about the junction
of the thin anterior third of the body with the more dilated
posterior portion. It seems highly probable that the manner in
which these parasitic Copepods can be so densely encrusted by
such organisms (other cases are referred to in the literature of
the subject), is directly related to the peculiar fact that after
fixation to their host they appear no longer to undergo ecdysis*.
Occurrence.—Kituta, Lake Tanganyika, 24.8.04. From the
soft region at the junction of the pelvic fins of a large Polypterus
congicus. One specimen.
White Nile. From the fleshy region at the junction of the
pair of pectoral fins of a Polypterus senegalus. Highteen speci-
mens, some of them incomplete, belonging to the collection of the
British Museum.
Fashoda, White Nile. Eight further specimens (one incomplete)
from the British Museum collection. The only particulars stated
are :—‘“ From Polypterus birchir, Fashoda.”
Tt is interesting to note that this species of Zerneocera has
been taken only on the Ganoid Polypterus, albeit on different
species of that genus. ,
3. LERNHOCERA TEMNOCEPHALA, sp.n. (PI. I. figs. 8 & 9.)
Description.—(Adult female.) Cephalic arms four, of unequal
size. The dorsal arms are long and stout, and fork distally in
a Y-shaped manner, terminating in rounded lobes. The ventral
arms are quite short, slender and simple, being bluntly pointed
at their ends. The body is almost straight, unsegmented and
cylindrical. The anterior third or less is moderately slender,
the body dilating gradually behind. Pre genital prominence
* Of. Jungersen, Mindeskrift for J. Steenstrup, xvi, Copenhagen, 1914, p. 6.
Proc, Zoou. Soc,—1914, No. LVI. 56
828 DR. W. A. CUNNINGTON ON PARASITIC
well marked, simple or slightly bilobed ; terminal portion of body
not upturned.
Total length of specimen, 10-0 mm.
From tip to tip of dorsal cephalic arms, 4-2 mm.
Remarks.—It is unfortunate that this species has to be
described from a single specimen. Although no egg-sacs are
present, it isalmost certainly adult, and there seems no reasonable
doubt that it represents a distinct form. This unique individual
has unluckily been badly preserved, and as a result the body
is nearly flattened towards its posterior end. It thus becomes
impossible to determine the real diameter of this part of the
body, and so to express the amount of dilatation which it has
undergone. Similarly, an artificial twist in the body makes it
virtually impossible to state the nature and degree of torsion,
From the shape of the cephalic arms, this form may be placed
not far from the oldest known European species, L. eyprinacea
and Z. esocina, from which it is nevertheless perfectly distin-
guishable. ‘The dorsal cephalic arms in this specimen, as will be
seen from fig. 9, are not quite bilaterally symmetrical, a condition
of things which is met with in the genus from time to time.
Occurrence.—The specimen belongs to the collection of the
British Museum. Particulars are given as follows :—‘ From
Barbus bynni, bought in Old Cairo fish bazaar—caught in the
Nile, Loat Coll., no. 26.”
3. General Remarks.
It is clear that, whatever may be the case for marine fishes,
the fishes of fresh water are relatively seldom the prey of parasitic
Eucopepoda under natural conditions. This is indicated by the
paucity of material in the collections under review, for the
specimens on which this paper is based come from eight indi-
viduals only. On the Third Tanganyika Expedition, of which
I can speak personally, very large numbers of fish were examined,
on only two of which were such parasites discovered. It is
instructive to compare with this the occasions on which Argu-
lidee—also external parasites—were obtained. They were taken
18 times in Lake Tanganyika, and in some cases on two or three
individuals of the same species at the same time, while with
lesser opportunities, they were found on three separate fish in
the Victoria Nyanza, as against no record at all of parasitic
Euecopepods. A study of the literature of the subject confirms
our conclusion. ‘There are, it is true, other families besides the
Lerneeide which are represented in fresh water, but they do not
constitute a very formidable assemblage, while the genus Lerne-
ocera, sole representative of its family, contains but a small
number of species. Of these species it is certainly true to say
that they are not very common, and in the majority of cases the
remark is equally true of the other parasitic forms. When
natural limits are transgressed, as they usually are in the stocking
of ponds and rivers with fish, there not infrequently occur
EUCOPEPODA FROM TANGANYIKA, 829
Serious epidemics due to vast numbers of parasites which have
multiplied under conditions abnormally favourable to them.
It is worth while to mention that a few other specimens
belonging to the British Museum have passed through my hands
without receiving any notice in the general text. This is because
they were too much damaged to permit of identification or
description, but they can be referred to the genus Lerneocera
almost with certainty, and as such are worthy of putting on
record. They were all taken on Nile fish, and came from three
separate individuals. One Specimen was taken on a Clarias
lazera, two more on another Clarias (2 lazera), while two-further
Specimens are labelled as follows :—“F rom Barbus bynani—
Damietta Nile, near Samannud. Loat Coll., no. 691.”
As regards the distribution of these African Species of Lerneidee,
it is to be noted that while one species has been found only
in Tanganyika and another only in the Nile, the third form
occurs in both. There is, then, no indication of that peculiar
nature and superior richness of fauna which in SO many cases
characterises the lake. At the same time, we know as yet so
extremely little of the distribution of these forms in Africa, that
it would be out of place to lay much stress on the facts which have
so far coine to light. It is, indeed, a fact that the Tanganyikan
species, Lerneocera diceracephala, is so fav peculiar that it ma y be
found necessary to establish a new genus for its reception, while
the two Nile species are much more typical members of the genus
Lerneocera. Further, if we deal with the parasitic Kucopepoda
as a whole, including the Ergasilidee, we still find that the fauna
of Tanganyika is exceptional and unusually rich in diverse forms.
Prof. Sars described * a new genus—FHrgasiloides—with three
endemic species from Lake Tanganyika, and from Lake N yasa,
only a single species of FErgasilus not further determined, but
which is probably the same as that recorded by Mrazek from the
Victoria Nyanza.
EXPLANATION OF THE PLATE.
(All figures by about 5.)
Lerneocera diceracephala.
Fig. 1. Ventral view.
2. Lateral view.
3. View of cephalic arms from above.
Lerneocera haplocephata.
Pig. 4. Ventral view.
5. Lateral view.
6. View of cephalic arms from above.
7. Lateral view of specimen infested with \ orticellids.
Lerneocera temnocephata.
Fig. 8. Ventral view.
9. View of cephalic arms from above.
* P.Z.S. 1909, p. 63.
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BRYOZOA FROM ZANZIBAR.
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BRYOZOA FROM ZANZIBAR. —
ON BRYOZOA FROM ZANZIBAR. 831
46. The Marine Fauna of British East Africa and Zanzibar,
. from Collections made by Cyril Crossland, M.A.,
B.Se., F.Z.S., in the Years 1901-1902. Bryozoa—
Cyclostomata, Ctenostomata, and Endoprocta. By
ARTHUR Wm. Waters, F.L.S., F.G.S.*
[Received May 19, 1914: Read June 9, 1914. |
(Plates L-IV.7 & Text-figure 1.)
INDEX.
Page
Geographical sDistnibuttommys ery ceeeneeessaet cesar snes ce Soe
Systematic:
Gristaenflacaaspelenreeenecee ee are Oe at Ee OOD
(OFROSOG): COREUMGAITy SDo Ts eosocasdadsbooosoucdsune cbobae esr Gon gbbor oodoaun ats AO)
Entalophora wasinensis, WOM. NOV. 22... ... eee eee eee eee esses. 840
Mimosella bigeminata, sp. 0. ...... 6.0 eee eee cece = 851
Zoobotryon pellucidum, rosette-plates ............ 6. eeeeeee eee eeeeee 849
The present paper continues the series on the collections made
by Mr. Crossland in the tropics. The first was on Vubucellaria,
published in the Journal of the Linnean Society, vol. xxx. 1907;
then two parts in the ‘ Reports of the Sudanese Red Sea,’
ibid. vol. xxx1. 1909, and vol. xxxi. 1910; and the Cheilostomata
from British Hast Africa and Zanzibar in the ‘ Proceedings’ of
the Zoological Society, 1913.
The bibliographical references are pretty full in order that, so
far as the tropics are concerned, they may be used as a con-
tinuation of Miss Jelly’s Catalogue.
It should not be forgotten that all the specimens dealt with in
this paper are from but a very slight depth (up to 10 faths.), so
that but few Cyclostomata would be expected, since they mostly
occur at considerable depths.
Points of Special Interest.
The ovicells of Hntalophora, and of Jdmonea radians Lam.,
have been studied full of embryos, and the occiostome has been
cut through. In Jdmonea radians the ovicells form lobes between
each series of zowcia. The ovicells must be more used in classi-
fication, and the primary zoccia and the early stages must be
examined.
The rosette-plates of Reichert in Zoobotryon, instead of having
a number of pores round a central pore, as described, have a
central pore and 8-9 cells round the pore. The mass of cells
in the neighbourhood of the pore are dealt with.
27 species or varieties are mentioned in this communication,
* Communicated by Cyrin Crossnanp, M.A,, B.Sc., F.Z.8.
+ For explanation of the Plates see p. 807.
852 |
854.
8352 MR. A. W. WATERS ON
and 76 were described in the previous one, making 103 in all;
of these 37 are known from the Atlantic, 22 are British, 31
Mediterranean, 37 are known from the Indian Ocean, 14 from
S. Africa, and 51 from Australia.
Table of Distribution.
== eed
Mediter-
yanean.
Australia.
British.
Red Sea.
Crisia denticulata (amk.) ..) +
.» elongata M.-Kd. | |
» sertularoides (Aud. & | | | |
NSPnv)} spews eee zl peut ate | |
» inflata, sp. u. aa
|
co COU REDITIEAT@I SYS Tilo soqnnooce |
+
+
+
++ | S. Africa.
ab
| Entalophora wasinensis, |
3. Tervia irregularis (Meneg.) .... + 2.5
. Idmonea milneana VOrb. ... + + + 2 7
7 | Amathia lendigera (li.) .........) + ) + | + 3 Sel ib.
' Mimosella bigeminata, sp. n....'
853 |
|| Buskia nitens Alder
|
KOS DNS | asecdeoueqcer +
Filisparsa tuhulosa (Busk) ... +B
oh
+
Pacific.
se radians (Lamk.) ......
© interjuncta MacG. ...
+++ 4+ 44+
+ + ++
: ices cae ee “Rs +
, Wistans Busk : io Meta
PS vidovici (eallayy bate te SRO ef Roscoff.
Zoobotryon pellucidum Ehy. .. | Isle of Pines,
Bowerbankia pustulosa(H.&S.) +. |
++
=
oa
+
WG DORTANGA RIS ememoncenocsocabaauee
Farrella atlantica Busk ...... +
Valkeria uva (li)... i ae
+
+ }
Sn can ete Pe ... | China Seas.
Cylindreecium giganteum By +
+ + .. ... | Queen Charlotte I.
Arctic, Q. Charlotte I.
- California.
+++:
' Pedicellina spinosa (Rob.) hee
Barentsia gracilis (Sars) ..... {hat
Loxosoma singulare Kef.
Lepralia poissonii Aud. ...... Neto laneesct tt eens Recetas
Japan.
Beania intermedia Hincks......) ... | ... | .. sone (ie
++
Levinsen * quotes Norman with regard to zoarial and zocecial
characters, who says: “ Why also in all instances is the ultimate
growth and form of the zoarium to he excluded from generic
characters among certain families of the Cheilostomata, and at
the same time to be recognised among the Cyclostomata and
Jtenostomata, and even some other groups of Cheilostomata ?”
Few characters have been available in the Cyclostomata, and
those mostly zoarial ones, so that the classification of the group 1s
still in a somewhat hopeless condition, whereas in the Cheilosto-
ae very many characters are now used, and the classification is
gradually becoming more natwal. It is therefore to the more
* Morph. and Syst. Studies on the Cheil. Bryozoa, p. 69 (19C9).
BRYOZOA FROM ZANZIBAR. 8353
highly differentiated Cheilostomata that we must at present look
for guidance, rather than to the simpler Cyclostomata.
There are, however, some characters in the Cyclostomata,
which when more used will help to show which forms are nearly
velated. Smitt showed that the ovicells furnished important
characters, and on several occasions I have pointed out the
importance of the ovicells, and of what I called the peristome of
the ovicellular opening, saying in 1888* that ‘“ the position and
nature of the opening is most important, eften more so than the
shape of the ovicell.” © Harmer has confirmed what I said, and in
several very valuable papers has gone into detail in some genera,
and has named the passage by which the larva escapes the
occiostome, while the external orifice is the oceciopore, which
terms are generally accepted, though unless new terms are
absolutely necessary they are always to be regretted, as every
branch of science 1s now overloaded with names. Also the size of
the embryo, and of the zocecial aperture, as well as the position
and nature of the closures, give specific or generic characters.
The primary zoecium and the growth of the younger part of
the zoarium must receive much more attention, in fact up to the
present in recent forms it has received hardly any: for example,
in Hntalophora proboscidea, E. deflera Couch, and Filisparsa tubu-
losa there is a Stomatopora-like growth often spreading for a
considerable distance over the supporting material, before the
zoarium becomes erect, whereas in what has been considered to
be Entalophora rugosa VOrb., from Naples, there is a Diastopora-
growth often covering a considerable piece of the stone or shell
upon which it grows before the erect cylindrical zoarium is
formed. It is elsewhere shown that other characters, namely
the lamina and ovicells, prove that it belongs to the Diastoporide.
To return to the more highly differentiated Cheilostomata : in
various genera the zoarium may be either adnate, erect, uni- or
bilaminate, or even articulated. For example, Lepralia, Schizo-
porella, Celleporidee, all occur adnate, erect, uni- and bilaminate,
while Zhalamoporella may be unilaminate, bilaminate, adnate or
erect and articulated ; Cellaride in recent forms are usually
articulate, except in the younger branches, but in fossils fre-
quently there is no articulation. Now in none of these cases is
there any material difference in the zoecium according to the
way in which the zoarium grows.
It is sometimes forgotten that all this only deals with the
position of the zoeecia, or, as Levinsen would say, the “‘autozooids,”
and this has proved of quite secondary value in classification ; but
this does not mean that characters furnished by the other
‘‘zooids” such as avicularia, stalks, stolons, radicles ete., which
also cause differences in the form of the zoarium, may not furnish
valuable characters.
No doubt the Catenicellide are derived from unarticulated
* “On some Ovicells of Cyclostomatous Bryozoa,’ Jour. Linn, Soc., Zool.
vol. xx. p. 276.
834 MR. A. W. WATERS ON
forms, but evidently this is a long way back, so that now
articulation is apparently a generic character; but there must
have been a time when closely related forms were in some cases
continuous, in others articulated, but now both zoarial and
zocecial characters indicate a group of Catenicellide. In Cellaridz
and Thalamoporella the articulation is more recent, and should
not be made a generic character.
Examining the Cyclostomata in the same way, we find to a
certain extent parallels with the Cheilostomata : Diastopora, or
perhaps we should say Diastoporide, occurs adnate, but also
bilaminate in such forms as D. intricaria, Mesenteripora; and
some forms now placed with Hntalophora, such as H. regularis
MacG., belong to this family. Even the D. obelia division, which
it has been proposed to raise to a genus Diplopora*, has adnate
forms, as well as the erect fossil Diastopora brendolensis Waters 7,
with tubules between the zoecia. A Stomatopora-like growth
may become erect, and too much importance has been attached
to whether a form is adnate or erect. There are adnate forms
with Heteroporidan structure, and some bilaminate as Pavospirua.
The ovicells together with the oceciostomes ete. are, as stated,
going to assist us to trace relationship, but to what extent we
cannot yet say, as our knowledge is in most families very in-
complete, often fragmentary or absent. However, although only
sufficient to show the direction in which work is wanted, it may
be useful to put together what I have gathered from my own
collection and from published accounts of recent forms.
In Crisiaz the ovicells of most species are known, and they with
the oceciostomes furnish most useful characters in determination.
In the species examined there are 8—9 tentacles.
In /dmonea, as at present understood, there are some important
and rather puzzling differences in the ovicells. There is, Firsv,
the /. radians, mentioned in this paper, also /. atlantica Forbes,
I. concava Reuss, and J. parasitica Busk, in which the oeeciostome
occurs on one side, usually on the second of the series enveloped
by the ovicell, and by the first or second zoewcium counting from
the median line; the tube curves over and turms downwards
(PI. IT. fig. 6, @.). In these the ovicell spreads across the anterior
surface.
Seconp. There may be merely an anterior inflation, usually
near to a bifureation, with a central ocwciostome, as /. interjuncta
MacG.
TurrD. I have a fragment of an /dmonea, probably australis
* This name cannot stand as it has already been used, see page 836.
+ Quart. Journ. Geol. Soc. vol. xlviti, p. 155, pl. iii, fig. 1 (1892).
~ Reuss (Foss. Polyparien des Wiener Tert. p. 99) described the Cyclostomatous
ovicells as Caelophyma, of which C. glabrum occurred on Crisia hérnesi (a Crisia),
Retepora disticha (apparently Idmonea), and R. cancellata (do. Idm.); and
Celophyma striatum on Hornera hippolithus. i
Hagenow (Bry. Maest. p. 105) described Calophyma levis on Truncatula repens
and T. truncata; C. constrictum on Idmonea tetrasticha (this is not Idmonea,
perhaps a worn Truneatula); and C. granulatwn on Idm. lichenoides. Gregory
does not quote Hagenow quite correctly.
BRYOZOA FROM ZANZIBAR. 835
MacG., in which the ovicell spreads over three or four series on
one side only of the median line.
FourtH. There is the very curious ovicell standing erect and
embracing the zocecia of a series, which I described * as occurring
in J. meneghini Hell., but from an examination of more material
T came to the conclusion that it was /. triforis Hell., and a
co-type has confirmed this conclusion, As described, there was
little to distinguish /. meneghina Hell. from J. triforis Hell.
except size, so that without a fair amount of material they were
not readily distinguished.
The zoceecial aperture of Jdmonea varies from 0:06—0°2 mm.
The so-called J. wregularis Meneghini has the ovicell dorsal,
and must be removed to Jervia. It has 13 tentacles.
In Tubulipora there is an anterior ovicell spreading among
many zocecia, with the oceciostome usually close up to a zoccial
tube. Zocecial aperture 0°07-0°18 mm. Tentacles 11-12. The
colony is, where attached, often provided with small projections
or even long multitubular radicles, so that the attachment is but
slight, whereas in Stomatopora it seems to be very close.
In Filisparsa there is an anterior ovicell. 14 tentacles.
In Entalophora few ovicells have been seen. In the proboscidea
group they are not very large, and are near to several zocecial
tubes without enclosing them. In wasinensis, the species
described in this paper, the ovicell is very long and contains
a considerable number of embryos. However, under /ntalophora
many species have been included which have a distinct lamina,
and in these the ovicells found are of the Diastopora type, and
suggest that they are erect cylindrical Diastopora. Entalophora
regularis MacG., and what I called #. rugosa dOrb., from
the Mediterranean, must be removed on this account from
Entalophora.
The zoecial aperture of Hntalophora is 0:07-0:19 mm. The
number of tentacles is 12-16.
Diastopora. The ovicell is vesicular, as an irregular or sub-
globular elevation, often involving many zoecia, and sometimes
situated tangentially to the colony. The ovicell may spread
internally with many arms, as in D. intricaria SmitttT. So far
as seen, the oceciostome of Diastopora is a small plain tube, usually
directed proximally. The zoccial aperture is 0:06—0°11 mm. %
The number of tentacles counted is 10-12. The ovicells of a very
considerable number of fossil Diastoporw have been figured, and
the genus was abundant in the Jurassic and Cretaceous periods,
and oceurs as Berenicea consimilis Lonsdale, in the Silurian (see
fig. in Bassler, ‘ Bry.-Fauna of the Rochester Shale,” p. 16, pl. v.
figs. 1-5, 1906).
* “QOvicells of Cyclost. Bryozoa,” Journ. Linn. Soc., Zool. vol. xx. p. 278, pl. xiv.
fig. 2 (1888).
+ Waters, “Bryozoa from Franz Josef Land,” Journ. Linn, Soc., Zool. vol. xxix.
p. 178, pl. xix. fig. 12 (1901).
t In some fossils the orifice is larger, see Canu.
836 MR. A; W. WATERS ON
Diastopora with tubules has been called Diplopora, a name
already used for a calcareous alga. MacGillivray made a genus
Diplopora, but finding the name occupied changed it to Diplo-
porella ; the name Diplopora has also been given for two or three
other things. The group, however, occurs incrusting and bila-
minate. The family Diastoporide will probably be found to be
more distinctly separated than any of the others.
Hornera. The ovicell is a large subglobular, dorsal or some-
what lateral chamber with large pits and a lateral oceciostome.
‘Tentacles, 9 in species examined. The ‘* Hornera eburnea”
Jull. & Calv. has a most curious anterior ovicell, but I do not
see why it is placed with Hornera; also the Hornera gravieri*
Calv. seems to have a somewhat similar ovicell. Zoccial
aperture of Hornera 0-04—0°12 mm.
Discotubigera (Defrancia). Tangential ovicell with oceciostome
near the distal border. One specimen has the ovicell the whole
way round the periphery. No doubt many species have been
placed under Lichenopora.
Stomatopora. Although many Stomatopore have heen figured
and described, but very few ovicells have been seen, and Smitt
said no species were known with ovicells. In 4. divergens
Waters t, the ovicell is a small round elevation on the anterior
surface. In S.(?) sp. from Plymouth the ovicell is at the end
of an erect branch as in Supercytis. In S. major Johns., the
anterior ovicell has a small plain tube for the oceciostome.
The S. compacta Norm. is Diastopora, and has the ovicells
raised in between the openings of a small number of zoacia, or
is tangential, with the occiostome as in Diastopora. Norman's
specimens examined are now in the British Museum,
Lichenopora. Ovicell central and spreading between the rays.
Oceciostome erect, plain or funnel-shaped; or a plain horizontal
tube low down near the edge of the ovicell. There may be many
oceciostomes, probably indicating several ovicells, just as a colony
of Diastopora may have a number of ovicells. Smitt mentions
eight occiostomes in L. verrucaria, and this must have been a
fine specimen, as I have never seen more than six. The zoecial
apertures in all recent species measured are about the same size
(from about 0-06—0-09 mm.).
Defrancia lucernaria Sarvs and Domopora stellata have the
ovicells in between the rays.
Frondipora has the ovicell across the anterior surface of a
branch, not much raised, and the occiostome, about 0:12 mm.
wide with the lower edge straight, also is but little raised, and is
not attached to a group of zocecia.
Flosculipora has the ovicell-wall uniting from neighbouring
zocecial bundles.
* Calvet, “ Bry. Cyclost. prov. des Camp. scient. accomp. p. S. A.S. le Prince de
Monaco a hord de Ja Princesse-Alice,” Bull. Inst. Ocean. No. 216, p. 7, fig. 6 (1911).
+ Krit. Fort. 1866, p. 414.
+ Expéd. Antarct. Belge, p. 89, pl. ix. fig. 6 (1904),
BRYOZOA FROM ZANZIBAR, 837
Heteropora., ‘The ovicell is unknown in recent species, but
Novak * figures one In a Cretaceous fossil. It is sac-like with a
small opening at one end. ‘Tentacles 14 in /. pelliculata Waters.
Supercytis has the ovicell at the end of the erect colony
spreading over the whole width. The Supercytis tubigera Busk, of
the ‘Challenger,’ is not related to Supercytis, nor is it correctly
described, as the series ave not uniserial but biserial. It looks
more like a Z’ubulipora.
Crisulipora. Oceciostome tube narrower than the zoccial tube,
without any terminal expansion, There are 10 tentacles.
CRISIA DENTICULATA (Lamarck), (PI. IV. fig. 5.)
Crisia denticulata Waters, “ Rep. Mar. Biol. of the Sudanese
Red Sea, Bry. pt. u. Cyclost. etc.,” Journ. Linn. Soc., Zool.
vol. xxxi, p. 232, pl. xxiv. figs. 1-3, pl. xxv. fig. 11 (1910); and
add :—
Osburn, “ Bry. of the Woods Hole Region,” cll, Bur.
Fisheries, vol. xxx. p. 216, pl. xviii. fig. 8 (1912); “ Bry. from
TLenaadi. etc.,’ Proe: Un. Sti Nat: Mus. vol. xliii. p- 276 (1912);
Guerin- Ganivet, “Bry. de la Région de Concarneau, ete.,” Trav.
Se. du Lab. de Zool. et de Phys. Mar. de Concarnean, ‘vol. ny
p- 19 (1912); “ Mission Arctiques: Bryozoaires,” Soc. d’Océan. du
Golfe de Gascoigne, p. 39 (1913); Osburn, “ Bry. of the Tortugas
Islands, Florida,” Pub, 182, Carnegie Inst. of Washington,
p. 185 (1914).
We see that the connections from the stolons to the zocecia, or
through the septa of the stolon, in Zooboiryon pellucidum Khy.
(p. 849) and other Ctenostomata are much more elaborate than
any description had indicated, a number of cells on each side
meeting those on the other, and to these groups of cells reach the
plasma network, which spreads through the stolon and the zoccia.
However, in Crisia and other Cyclostomata, I have not found a
plasma network spreading all through the zocecium, as we know
it in Cheilostomata and Ctenostomata, but near the base below
the ceecum there is a small number of threads. The polypide so
nearly fills up the zoecial tube, that there does not seem to be
room for much network of plasma.
In Crisia and cther Cyclostomata, the number of connecting
pores is very considerable, being situated generally all along the
surfaces, and, as a rule, the zoccium at its proximal end is
connected with the zoccia on the two sides, in such a way that
it seems impossible to speak of a new zoecium having arisen
from any one older zocwcium (PI. 1V. fig. 6). There are “but few
cellsin contact with the pore, in this case apparently two on each
side (PI. IV. fig. 5).
Loc. Add: Arctic; Atlantic, Cape Verde Islands. Wasin, Brit.
K. Africa, 10 fath. (601, 516).
* “Bry. der béhmischen Kreideformation,” Denks. der Math.-Naturw. Cl. K.
Akad. Wien, vol. xxxvii. pl. yini. figs. 80, 31 (1877)
838 MR. A. W. WATERS ON
CrisiA ELONGATA Milne-Edwards. (Pl. I. figs. 3, 4; Pl. IV.
fig. 6.)
Orisia elongata Milne-Edwards, ‘“ Mém. sur les Crisies, les
Hornéres,” Ann. des Sciences Nat. ser. 2, vol. ix. p. 10, pl. vu.
fig. 2 (1838); ? Busk, Brit. Mus. Cat. Cyclost. p. 5, pl. iv. figs. 5,
6 (1875); ¢Busk, Chall. Exp. Zool. vol. xvii. p. 5, pl. i. fig. 3
(1886).
The specimens from Wasin are without any doubt the species
described by Milne-Edwards, even though he says * plus gréle”
than C., denticulata Lamk., which is not the case. Busk, in his
Museum Catalogue, speaks of the zocecia being much produced,
whereas this is never the case in my specimens, nor does Milne-
Edwards show it, or even Busk himself in his figures.
The lateral branch, of which there is one, and only one, to each
joint starts from near the end of a joint, after about the 6th-10th
zocecia on the one side. The last zocwcial tube is continued free,
as is the case to a certain extent with the last zocecium on the
other side. The number of zoccia is uneven, and in some of the
terminal nodes as many as 13 pairs of zocecia have been counted.
There is the small mark below the oral aperture, as in C. denti-
culata, showing its relationship. The older chitinous joints are
black, the younger ones are light. No ovicells are known.
The surface has numerous pores, and | do not understand Busk
speaking of it as granular. The closure is slightly raised in the
centre, and near this there are one or two pores.
The zoarium is about 0°3 mm. wide; the distance from zocecium
to zocecium, on the same side, is about 0°25 mm.,and the aperture
of the zocecia is about 0°07 mim.
Since I wrote my Naples paper, I have been able to examine
better specimens of what I took to be C. elongata, which correspond
most nearly with C. cribraria Stimpson, as re-described by
Osburn *.
In the Naples specimens, the fresh internode arises after the
2nd, 3rd, or 4th zoccium of the one side, and another branch
arises from after about the 6th zocecium on the other side; no
ovicells were found, though some are forming at the very end of
the branch, and the zowcia are about 0°45 mim. apart, which is
about the same as in C. ramosa H., with which it is allied, but
the chitinous joints are light. What I called var. angustata,
I now consider is C. ramosa Harm.
The Algoa Bay specimen, so described in the British Museum
Catalogue, is probably elongata, and in this specimen the fresh
branches are always high in the internode. The ‘ Challenger’
specimen named elongata I do not think is this species, and it
has branches both high and low in the internodes, more as in
C. ramosa. The specimens called elongata by Norman from
Madeira are more like C. rwmosa.
Loc. Red Sea ? (I.-Hd.); Algoa Bay? Wasin, Brit. E. Africa,
10 fath. (501), collected by Crossland.
* “ Bryozoa of the Woods Hole Region,” Bull. Bur. of Visheries, yol. xxx. p. 215,
pl. xviii. fig. 7 (1910).
BRYOZOA FROM ZANZIBAR. 839
.
CRISIA SERTULAROIDES (Audouin & Savigny) *. (PI. I. figs. 5, 6. )
Proboscina sertularoides Aud., “ Descrip. de ’Kgypte,’’ Hist.
nat. p. 236; Savigny, pl. vi. fig. 6.
Crisia recurva Heller, “ Bry. des Adriat. Meeres,” Verh. d. K.-K.
zool.-bot. Gesellsch. Wien, vol. xvii. p. 118, pl. vi. figs. 3, 4 (1867).
Zoaria of rather straggling growth, apparently about 20 mm.
high; internodes short, with 7-15 zocecia, though usually 7-9,
with light joints, and the branches grow from above the first
zoceclum on that side, with sometimes another branch on the
other side, near the distal end of the node. Branches not very
wide (about 0°15 mm.); zoccia directed forwards with only a
short part free, the distance from zocecium to zowcium is about
0-21 mm., the zoccial aperture is 0°06 mm.
The ovicells are very wide and large, irregular globular, placed
to one side, and the oceciostome is funnel-shaped.
The growth is like that of C. eburnea L., but the branches arise
higher up after the first or second zocecium, and in nodes where
there is a branch the number of zoccia is uneven. It is also
much like C. aeropora Busk, though the denticle by the aperture
occurs but very rarely; from C. ramosa Harm., it differs in
having shorter internodes and in having the zoecia much nearer
together. Although Heller’s figures and description are unsatis-
factory, it does not seem that there can be any doubt as to the
identity of the species.
A specimen from Ras Osowamembe (504) has the zocecia less
spread out, the joints black in the older parts, and toe zoccia
about 0°28 mm. apart; the ovicells are large and irregular over
the whole width of the zoarium. ‘The difference may only be
local, and it seems advisable to consider it a variety.
Loe, Adriatic (Heller). Wasin, Brit. KE. Africa, 10 fath, (500) ;
Chuaka, Zanzibar, 3 fath. (506), collected by Crossland.
CRISIA INFLATA, sp.n. (PI. I. figs. 1, 2.)
This is one of the most delicate species of Crisia, and is some-
what like C. geniculata M.-Ed., but there are two alternate zocecia
in a joint, the new node rising from the end of the last. The
nodes of (C. geniculata are ee to three times as long, and the
zocecial apertures of geniculata are about 0-075 mm., whereas i in
inflata they are only about 0:04-0:05 mm. The very light
corneous tubes of the joint are, when examined with a high
power, found to be marked with fine longitudinal lines.
The ovicell is suberect and is only an inflation of the zocecial
tube, being the simplest ovicell known. From this character the
specific name is chosen. The oceciostome is dorsal, so that it
cannot be seen from the front, and is a small plain tube curved
over, with a circular aperture 0:02-0:03 mm. The ovicell is not
* (The parentheses around the names of authors placed after scientific names in
this paper are used in accordance with Article 23 of the International Rules of
Nomenclature (Proc. 7th Int. Cong., Boston 1907, p. 44 (1912))—Enprtor. |
840 MR. A. W. WATERS ON
elongate like the ovicell of geniculata, as figured by Busk and
Harmer, and also as a specimen in my collection, Harmer
considers that C. geniculata and C. cornuta must be separated,
chiefly on account of differences in the ovicell, and although the
present species is also allied there seem to be sufficient reasons
for its separation. The C. crisidiodes Ort. has 2-3 zowcia to each
internode, with three-jointed sete.
Los. Wasin, Brit. E. Africa, 13 fath. (500), only one colony.
Crista CIRCINATA, sp.n. (PI. I. figs. 7-9.)
The zoarium divides into two main branches, and on each of
these the fresh branches are mostly given off from the one side,
and in the lower part of the zoarium the fresh branch arises at
about the second zocecium, while in the younger part the branches
start from about the fourth. The joints are light. The distance
from zocecium to zocecium is about 0°27 mm., and the round
zowcial aperture is about 0-08 mm. ‘There are fewer pores on
the zocecia than in most Crisiw, and the zocecia extend free for a
considerable distance.
The ovicells, occurring just after a bifurcation, are large and
erect, with the curved ocwciostome on the distal or dorsal surface
of the ovicell, with the oceciopore only 0°05 mm. diameter.
The ovicell of Crisia may appear to be central as in C. ramosa
Harm., C. fistulosa Hell., etc., and is then long and pear-shaped,
or it may appear to be at one side, and may be shorter, when
we call it pomiform. There are others in which the ovicell is
free, not being attached by its surface, and with the oceciostome
on its dorsal surface instead of being directed forwards. Free
ovicells occur in C. edwardsiuena VOrb., C. biciliata MacG.,
CO. howensis Maplestone.
IT have felt much hesitation as to whether the form described
is the Crisia cuneata Maplestone *, and it also has many points
of resemblance with C. cylindrica Busk, but a different ovicell is
figured. The Museum specimens (528, 853) of eylindrica have
no ovicells. i
Loc. Ras Osowamembe, Zanzibar Channel, 10 fath., only one
specimen (504); Prison Island, Zanzibar Channel, 8 fath. (505) ;
Brit. E. Africa, 10 fath. (520): collected by Crossland.
ENTALOPHORA WASINENSIS, nom. nov. (Pl. II. figs. 1-4, 9;
Text-fig. 1.)
Entalophora deflexa Smitt, “ Floridan Bryozoa,” pt. i. p. 11,
pl. v. figs. 28-30 (1872).
As this does not seem to be the same as the small delicate
British species known as deflewa Couch, I have given it another
name.
The specimens from Zanzibar are buried in sponge, which has
* “Word Howe Island Polyzoa,’ Proc. Roy. Soc. Vict. vol. xvii. n.s., p. 390,
pl. xxix. fig. 12 (1904). i
BRYOZOA FROM ZANZIBAR, 841
grown over them. The zoarium is about 25 mm. high, dividing
into many new branches at rather an acute angle. There are
many zocecia, with long tubes all round the zoarium, with the
oral aperture 0-09 mm.—O'l mm. There are 12 tentacles, and
transverse sections show about 15-16 zoecia.
The base of this species has not been seen, but #. proboscidea
grows from a multiserial Stomatopora-like base, which often
spreads for some distance before the erect growth commences.
EL. defleca also starts from a Stomatopora base which is
Text-figure 1.
Entalophora wasinensis. * about 4.
principally uniserial, though in places it may be biserial. The
zocecia of this species, HZ. elegans Novm., and £. deflexa Couch,
have the aperture about the same size and are allied ; and as the
zoarial growth of H. elegans Norman ™* is similar, it is doubtful
* * Polyzoa of Madeira,” Journ. Linn. Soc., Zool. vol. xxx. p. 281, pl. xxxv. figs. 4, 5
(1909).
842 MR. A. W. WATERS ON
whether the greater projection of the zoccial tubes may not
depend on local conditions.
Only few ovicells have been found: one is small at the side
just above a bifurcation, and has an oceciostome with a narrow
opening and a somewhat triangular plate in front of it. The
other ovicells have the occiostomes like those in the first, and
spread round several zocecia without including any—that is, none
pass through it. This structure was shown by Smitt (fig. 30).
Very few ovicells of recent Entalophora have been described.
Busk says of 2. delicatula B. “ocecium tumid,” but it has never
been figured. In the ‘Challenger’ Report Busk, on pl. iv.
fig. 1b, figures an irregular zocecium as an “ ovicell dilation,” but
from an examination of the specimen I think this is a mistake,
as there is no sign of more numerous perforations, which
we seem always to find on the ovicells of these groups of
Cyclostomata.
In my collection there are ovicells on 2. regularis MacG. and
FE. intricaria Busk*, but these and some other species, among
which what I considered was 2. rugosa VOrb., from Naples, will
probably have to be removed from Hutalophora, as sections show
a distinct lamina, and the ovicells are wide with the zocecial
tubes passing through, reminding us of the ovicells of Diastopora,
with which they are closely allied, but whether they must be
ealled Diastopora or Bidiastopora need not now be considered.
Among fossils also, no doubt many must be removed from
Entalophora to Diastoporide. D’Orbigny and others have also
placed under Hntalophora many species now remoyed to
Meliceritites. .
The ovicells of fossil Hntalophera (Spiropora) annulosa Mich.
are figured by Canu‘, and are fairly similar to the ovicells of the
present species. Canu does not figure the zocecia spiral or
regular, therefore why does he call it Sprropora ?
Loc. Flovida (Smitt). Wasin, Brit. E. Africa, 10 fath. (501)
(507); Ras Osowamembe, Zanzibar Channel, 10 fath. (504):
collected by Crossland.
FrnispARSA TUBULOSA (Busk).
Hornera violacea var. tubulosa Busk, Cat. Mar. Poly. pt. iii.
p- 19, pl. xviii. fig. 4: for synonyms see Waters, “ Austral. Bry.”,
Ann. Mag. Nat. Hist. ser. 5, vol. xx. p. 257 (1887); and “ Rep.
Mar. Biol. of Sudanese Red Sea, Bryozoa,” Journ. Linn. Soe., Zool.
vol. xxxi. p. 235, pl. xxv. figs. 16, 17 (1910); and add: Seguenza,
Form. Terz. pp. 297, 372 (1879); De Stefani, “ Jejo Montalto e
* E. intricaria Busk has the rays or spines on the outside of the zoccial tubes
which pass through the ovicells. The lamina is not seen in all stages, and in the
section which I figured, Q. Journ. Geol. Soc. vol. xliii., pl. xviii. fig. 5, none is seen
though in other pieces it is quite distinct. j
+“ Btudes sur les ovicells des Brvoz. du Bathonien d’Occaignes,” Bull. de la Soc.
Géol. de France, 3™¢ sér. vol. xxvi. p. 282, figs. 19, 20 (1898).
BRYOZOA FROM ZANZIBAR. 843
Capo Vat.”, Mem. R. Accad. d. Lincei, vol. xviii. p. 208 (1882),
Neviani, “ Bri. Neog. delle Calabrie,” Pal. Ital. vol. vi. p. 234
(1900).
As Busk made a mistake about numbering his figures, it
is difficult to understand what he meant, but in spite of this
it seems advisable to retain his name. Apparently the type
violacea also should be Filisparsa, and in the Norman collection
specimens so named, from Flord, have none of the appearance of
Hornera, but look like Filisparsa, having anterior ovicells which
are broken down. There is among these specimens one piece of
Hornera, perhaps lichenoides, evidently misplaced.
The specimens from Wasin have the oral aperture about
0-15 mm., corresponding in this respect with specimens from
Naples and Australia, and here also the closure of the zoccial
tube has a number of perforations similar in appearance to those
of the zowcia. There are 14 tentacles. The ovicell spreads
over the front, and the oceciostome, directed somewhat backwards,
is about the width of the zoccial aperture, but in the longi-
tudinal axis is only about 0-°O07mm. There can scarcely be said
to be a funnel, although there is an irregular expansion which
frequently curves over.
The expansion and funnels of the occiostomes of the Cyclosto-
mata are often very variable, so that too much weight must not
be attached to their measurements; also some ovicells may be
found with and others without funnels.
There are some specimens from Zanzibar, apparently of this
species, which in younger parts just touch the support at intervals,
though there are no definite rows of dorsal projections as in
Tubulipora pulchra MacG.; in the older parts there are strong
calcareous radicles, sometimes formed of only one tube, but more
frequently of two or three, which may divide at the end to form
claspers. Jdmonea pedata Norman has still larger radicles the
width of the zoarium, often formed by five or six tubes.
A species from Naples, which I have considered to be the
Tubulipora incrassata d’Orb. as more fully described and figured
by Smitt, has zoecia much about the same size, and standing up
in the same way; but the zoarium spreads out fan-shaped, instead
of being unattached for a considerable distance, and is continuous
about the same width. ‘The ovicell is situated as in 7’. tubulosa,
with the oceciostome about the same size as the zoccial tubes,
sometimes ending off straight, at other times with a funnel.
Loc. Naples (W.); Victoria (MacG.); Holborn Island, Queens-
land, 20 fath.; North Atlantic (4.). Ras Osowamembe, Zanzibar
Channel, 10 fath. (504), collected by Crossland.
Fossil. Tertiary : Rhodes, Sicily, ete.
TERVIA IRREGULARIS (Meneghini). (Pl. IV. fig. 8.)
Idmonea wregularis Meneghini, “ Polipi della fam. dei Tubul.
finora osserv. nell’ Adriatico,” Nuovi Sagei del Accad. di Scienze,
Proc. Zoot. Soc.—1914, No. LVII. o7
844 MR. A. W. WATERS ON
Padova, vol. vi. p. 12 (1844). For synonyms see Miss Jelly’s
Catalogue and add :—
Filisparsa irregularis Waters, ‘“Ovicells of Cyclost. Bry.,” Journ.
Linn. Soc., Zool. vol. xx. p. 279, pl). xiv. figs. 5, 6 (1888); Norman,
“ Poly. of Madeira,” Journ. Linn. Soc., Zool. vol. xxx. p. 279,
pl. xxxiv. figs. 1-3 (1909).
Tervia folini Calvet, “ Rech. de la Camp. du ‘ Caudan,’” Ann.
de l’Univ. de Lyon, p. 265, pl. vil. figs. 1-3 (1896).
Tervia irregularis, Jull. & Calv. “ Bry. de VHirondelle,” p. 114,
& p. 157, pl. xiv. fig. 7 (1903).
The proximal part of the zoarium has the zoecia irregularly
placed as in Filisparsa; then later, usually after the first
branching, there are distinct series on each side, often with isolated
zocecia in the median line. The very earliest part, namely the
primary, is like that of Stomatopora, being about the same size as
ordinary zocciaand but very slightly expanded at the proximal
end, whereas in Jxubulipora and many other Cyclostomata there
is a large disk. This disk I have figured mm 7’ubelipora pulchra,
and it has also been figured by Barrois, Robertson and others.
Inside the zocecial tube, about the position where it becomes
erect, there is on the proximal side a shallow comb-like process
(Pl. IV. fig. 8). This is where the closure takes place. There
are combs in some Membraniporide, and spimous processes in
many Cyclostomata, but I have not found a similar comb in any
other Cyclostomata and it does not oceur in Filisparsa tubulosa.
Loc. Adriatic; Naples, 40 fath. ; Genoa: Bay of Biscay :* Azores,
450 fath. (J. & C.); Madeira (V.); off Cape Blanco, West Africa,
235 met. (J. & C.); Australia, Wasin, Brit. E. Africa, 10 fath.
(507), collected by Crossland.
TDMONEA MILNEANA d Orbigny.
For synonyms and localities, see Waters, “Bry. from near Cape
Horn,” Journ. Linn. Soe., Zool. vol. xxix. p. 249 (1904).
From Ras Osowamembe, Zanzibar Channel, 10 fath. (504),
collected by Crossland.
IDMONEA RADIANS (Lamarck). (PI. IT. figs. 6, 7, 8, 10.)
For synonyms see Miss Jelly’s Catalogue and add :—
Dollman, W. P., Journ. Roy. Micr. Soc. pl. viii. 1906, photo-
graph only; Philipps, E. G., “ Polyzoa collected by Dr. Willey,”
Willey’s Zool. Results, pt. iv. p. 449 (1899); MacGillivray,
‘“Monog. of the Tert. Polyzoa of Victoria,” Trans, Roy. Soe. Vict.
vol. iv. p. 121, pl. xvi. fig. 18 (1895).
Besides the more usual form with short branches (fig. 8a) there
are from both localities specimens with long ones (fig. 8) having
similar subparallel branches. MacGillivray has referred to a
larger form, and Busk called it var. erecta. In the smaller
BRYOZOA FROM ZANZIBAR. 845
form as a rule there are two zoccia to a series, though there
may be three, especially near the growing ends, while the larger
form has three zocecia or occasionally four. A range of inter-
mediate sizes have been met with.
The ovicells are anterior, near a bifurcation, or in other
positions, even often half-way between two bifurcations; large
pores or pits occur onthe surface. The oceciostome is on one side,
most frequently by the first zovecium of the second series involved
(fig. 6), though sometimes it occurs by the third series, but never
more than one occiostome has been seen on an oviceil, and the
tubular oceciostome turns over and downwards, resembling in
position and form the oceciostomes of J. atlantica F., [. concava
Rss., and /. parasitica Busk.
The large lateral plates on the ovicell described by me* as
occurring in Torres Straits and Australian specimens, are not
seen in those from Zanzibar, though from the localities previously
mentioned they are very distinct, and are also seen in the
‘Challenger’ specimens from Tongatabu, and in the British
Museum specimens from Cape Capricorn, where however they
are elongate rather than round. There are no frontal ridges on
the ovicell as described by MacGillivray. The series are about
0-26 mm. apart, which is much closer together than in any other
species measured. J. atlantica F. is 0-6-1 mm.; J. australis
MacG. 0°75 mm.; J. concava Reuss 0°5 mm.; L. pedata Norm.
0-45 mm.; J. tumida Sm. O17 mm. The zoccial aperture is about
0-06 mm., whereas in J. atlantica it is 0°15 mm.; in J. milneana
0-:16-0-2 mm. There are 8 tentacles.
The ovicell consists of several lobes, there being on both sides
one between each two series of zocecia, so that in a mature ovicell
there are usually six such lobes. The embryos are small; the
mature ones may be 071 mm. across or even a trifle larger, but the
majority are smaller. There is not much difference from the
embryos of Hntalophora wasinensis nov., though a little larger,
but they are smaller than those of Diastopora intricaria Sm.,
0-12 mm.
Ostroumoff? says the larvee of the Cyclostomata vary in size
downwards from Hornera which is 0:48 mm., through Tubulipora,
Frondipora, Lichenopora, to Crisia, which last is only 0-07 mm.
In sections I have found Hornera lichenoides 0'4 mm. and various
species of Crisia from 0:07-0°1. The range in the Cheilostomata
is somewhat similar, as Diplodymoides is 0°08, and Systenopora
0°37 mm.
This is no doubt Crisina hochstetteriana Stoliczka t, but the
* “ Bry. from New South Wales” etc., Ann. Mag. Nat. Hist., ser. 5, vol. xx.
p- 254, pl. vi. figs. 27,28 (1887).
+ “Zur Entwickelungsgeschichte der Cyclost. Seebryozoen,” Mitt. Zool. Stat. zu
Neapel, vol. vii. p. 180 (1887).
{ “ Foss. Bry. der Grakei Bay bei Auckland,” Novara Expedition, p. 118, pl. xviii.
fig. 3 (1864). :
7 57*
846 MR. A. W. WATERS ON
I. hochstetteriana of MacGillivray * is clearly Hornera? fissurata,
Busk.
The name Jdmoneat is used in the sense it has been used for
a long time, although recognising that im some cases it is difficult
to ball distinctions Tatween Idmonea and Tubulipora, and that it
is possible they may have to be merged.
But Tubulipora spreads out continuously, whereas Jdmonee
continues of the same width; also 7’wbuliporw has a more or less
central ovicell with a central occiostome, while Jdmonea usually
has the oceciostome near to the series at one side. Perhaps
further study of the primaries and early growth, as well as the
ovicells, may establish the position, but for the present no harm
is doue by waiting until the whole of the group is better under-
stood.
Loc. New Zealand, Wanganui, ete.; Tongatabu, 18 fath. ; Hon-
olulu, 20-40 fath.; Victoria ; Port Jackson (H.), Adelaide (4. Wy
Sydney ; Port Stephens, 5-6 ‘fath.; Green Point, 8 fath.; Darnley
Island, Torres Straits, 10-30 ea ; Cape Capricorn, B.M. coll. ;
Lifu (Ph.); Port Elizabeth (in Miss Jelly’s collection). Both the
small and the var. erecta forms from Ras Osowamembe, Zanzibar
Channel, 10 fath. (504), and Prison Island, Zanzibar Channel,
8 fath. (505), collected by Crossland.
Fossil. Orakei Bay, N. Z.; Mount Gambier, S. Aust.; Bairns-
dale, Gippsland.
IpMONEA INTERJUNCTA MacGillivray. (PI. IT. fig. 5.)
Idmonea interjuncta MacG. Trans. Roy. Soc. Vict. vol. xxii.
p- 137, 10 (sep.) (1886); Waters, “ Austral. Bry.,” Ann. Mag.
Nat. Hist. ser. 5, vol. xx. p. 256, pl. vi. fig. 29 (1887).
Idmonea pedleyi Haswell, ‘‘ Cyclos. Polyzoa of Port Jackson,”
Proc. Linn Soc. N.S. Wales, vol. iv. p. 351 (1880).
Idmonea pulcherrima Kirkpatrick, ‘“‘ Hyd. and Poly. from the
China Sea,” Ann. Mag. Nat. Hist. ser. 6, vol. v. p. 22, pl. iv
fiz. 6 (1890).
The zocecial aperture is about 0°12 mm., and the oceciostome is
the same'size.
Some specimens from Ras Osowamembe have faint ridges on
the dorsal surface, and by the side of these ridges are rows of
pores, transverse to the zoarium. Lines of pores, though not as
marked, are also found on Jdmonea milneana d’Orb., a species in
many respects similar, but milneana is a larger species having the
zocecial aperture larger.
Loc. Port Phillip Heads (MacG.); Green Point, Port Jackson,
8 fath. (Waters). Ras Osowamembe, Zanzibar Channel, 10 fath.
* “Tert. Polyzoa of ee as Trans. Roy. Soc. Vict. vol. iv. p. 120, pl. xvi.
figs. 12-16 (1895).
“+ See my re-deseription “ On some Oviceils of Cycl. Bry.”, Jonrn. Linn. Soe., Zool.
vol. xx. p. 275, pl. xiv. figs. 1, 3, 4, 7 (1888).
ose Harmer, “ Devel. of Tubulipora,” Quart. Journ. Micr. Sc. vol. xli. n.s. p. 88
BRYOZOA FROM ZANZIBAR. 847
(504); Prison Island, Zanzibar Channel, 8 fath. (505), collected
by Crossland.
AMATHIA LENDIGERA (Linneus). (Pl. IV. figs. 3, 4.)
For synonyms see Miss Jelly’s Catalogue and add :—
Amathia lendigera (L.), MacGillivray, “On the Australian
Species of Amathia,” Proc. Roy. Sec. Vict. vol. vii. p. 185, pl. B.
fig. 1 (1894): Calvet, “‘ Bry. Ectoproctes,” pl. vill. figs. 19, 20,
pl. xiii. figs. 138, 21 (1900); “ Bry. Mar. des Cotes de Corse,” p. 46
(1902); “ Bry. Mar. de la Région de Cette,” p. 90 (1902); Jullien
& Calvet, “ Bry. de |’Hirondelle,” p. 31 (1903) ; Guerin-Ganivet,
“Bry. de la Rég. de Concarneau,” Tr. Se. du Lab. de Zool. et de
Phys. Mar. de Concarneau, vol. iv. p. 23 (1912).
The stems of the specimens from Chuaka are about 0-07 mm.
in diameter. This species differs from A. videvei Hell. in not
having the zoecia spiral, also in the branching being more or
less at right angles to the main branch (fig. 4), whereas in most
Amathia the branches divide equally in both directions. Hincks
has figured the branching of lendigera.
Some sections of material from Swanage cut across the rosette-
plate show the semicircle of cells, with nuclei at the end of the
cell, directed to the pore; also the round mesenchym-cells with
round nuclei are separated from the wall of the stolon, and are
seen to pass over the mound of cells; up to this mound come the
funicular threads with their elongate nuclei (fig. 3). The struc-
ture of the cells near the rosette is very similar to that described
in Zoobotryon (see page 850).
Loc. British, French and Belgian coasts, Mediterranean, Adri-
atic ; Corsica (40-60 met.) (Calvet); Azores (J. &@ C.); Western
Port, Victoria (acG.). Chuaka, Zanzibar shore (521, 523),
collected by Crossland.
AMATHIA SEMICONVOLUTA (Lamouroux).
Amathia semiconvoluta Lamx. Encycl. Méth., Zoophytes, p. 44
(1824); Heller, “‘ Bry. Adriat.,” Verh. der K.I. zool.-bot.
Ges. Wien, vol. xvii. p. 127, pl. v. figs. 1, 2 (1867); Calvet, Bry.
Kctoproctes, pl. vii. figs. 8, 9, pl. viil. figs. 16, 18 (1900); “ Bry.
Mar. de la Région de Cette,” Tr. Inst. de Zool. de |’Univ. de
Montpellier, ser. 2, mém. 11, p. 89 (1902); Waters, Journ. Linn.
Soc., Zool. vol. xxxi. pl. xxiv. fig. 6 (1910).
Serialaria seniconvoluta Lamk. Hist. Nat. d’Anim. sans vert.,
ed. ii. vol. ii. p. 171 (1836); d’Orbigny, Pal. Frang. vol. v. p. 595
(1850 -52).
The yellowish thick chitin-stem is about 0°25 mm. diameter.
Near the proximal end of each internode there is a clear oval
spot, which is for the attachment of a radicle, yet although this
mark for the attachment occurs in all internodes, radicles have
only been seen in a very few cases. A similar mark for radicles
oceurs in A, obligqua MacG., and also in A. brasiliensis Busk, of
848 MR. A. W. WATERS ON
the ‘Challenger,’ which no doubt is really A. semzconvoluta, but
the character has not been found in any other species examined.
Loc. Mediterranean ; Naples (A. W. IW. coll.) ; Adriatic (Heller).
Wasin, Brit. E. Africa, 10 fath. (501), collected by Crossland.
AMATHIA DISTANS Busk.
Amathia distans Busk, Chall. Exp., Zool. vol. xvii. pt. 50. p. 33,
pl. vii. fig. 1 (1886): MacGillivray, “On some 8. Australian
Polyzoa,” Trans. Roy. Soc. 8. Australia, vol. xii. p. 30 (1889);
“On the Austral. Sp. of Amathia,” Proc. Roy. Soc. Victoria, vol.
vii. p. 134, pl. C. fig. 3 (1894); Waters, ‘ Mar. Biol. of the Sudanese
Red Sea, Bryozoa,” Journ, Linn. Soc., Zool. vol. xxxi. p. 243,
pl. xxiv. fig. 7 (1910).
The A. distans B., A. tortwosa B., A. semiconvoluta Lamx., and
A, vidovici Hell. form an incomplete spiral round the stem, only
occupying part of the internode. The stem of A. distans is thin,
measuring 0:07 mm. diameter in most of the present specimens.
The ‘ Challenger’ specimens which I measured have the diameter
about O'l mm. I seem to have measured some abnormal piece
from Zanzibar which was stouter. From bifurcation to bifurcation
is about 2 mm. long. There are 8 tentacles.
Loc. Off Bahia, 10-20 fath. (B.); South Australia (MacG.) ;
New South Wales (4. W.W. coll.). Zanzibar town, shore, marked
“pink Polyzoum” (527), collected by Crossland.
AMATHIA yipovici (Heller). (Pl. IV. figs. 1, 2.)
Valkeria vidovici Heller, “ Die Bry. des Adriat. Meeres,” Verh.
der K.K. zool.-bot. Ges. Wien, vol. xvii. p. 128, pl. v. figs. 3, 4
(1867).
Vesicularia dichotoma Verrill, ‘Invert. animals of Vineyard
Sound,” Rep. Comm. of Fish and Fisheries for 1871-2, p. 874.
Amathia lendigera Busk, Chall. Exp., Zool. vol. xvii. p. 33 (1886).
Amathia dichotoma Osburn (Verrill), “ Bry. of the Woods Hole
Region,” Bull. Bur. of Fish. vol. xxx. p. 254, pl. xxix. figs. 81, 81 a
(1912).
The zocecia at a bifurcation are in short biserial clusters, spirally
arranged, encircling the stolon. There is sometimes a small
cluster between a bifurcation as shown by Heller, whose figure
was evidently froma dried specimen and is not entirely satis-
factory.
The branches are about 0-2 mm. in diameter. There are 8
tentacles, which is the same as in A. lendigera L., A. senricon-
voluta Lamx., A. brongniartii Kirkp., and A. distans B. ‘he
gizzard is about 0-05—0-06 mm. diameter.
The connecting cells on the two sides of the rosette-plates are
fairly similar to those of Zoobotryon pellucidum Hhbr., radiating on
both sides from the pore, and the cells near this pore stain more
darkly than the others (fig. 2).
Loc. Adriatic (Heller); Genoa (Waters coll.) ; Roscoff, sent to
BRYOZOA FROM ZANZIBAR. 849
me by Jullien as A. semiconvoluta; Bermuda, 30 fath. (Chall.) ;
Great Egg Harbour, N. J.; Long Island Sound (Verrill) ; Vine-
yard Haven, Edgartown, Woods Hole, Nantucket (Osburn).
Wasin, Brit. E. Africa, 10 fath. (500) (501), collected by Cross-
land.
ZOOBOTRYON PELLUCIDUM Ehrenberg. (Pl. III. figs. 4-12;
PAP Ve tiestl2-)
For synonyms see Waters, ‘‘ Mar. Biol. of the Sudanese Red Sea,
Bryozoa,” Journ. Linn. Soc., Zool. vel. xxxi. p. 243, pl. iv. figs. 12,
15 (1910), and add 22 Oshur n, “Bry. of the Tortugas Toland!
Florida,” Pub. 182, Carnegie Inst. of W ashington, p. 218 (1914).
The appearance of the specimens from Chases Bay, at first
sight, suggest specific separation from Z. pellucidum, as on the
whole length of the long internodes there are two distinct, wide
series of zocecia closely crowded, but with vacant longitudinal spaces
between the series. In the Naples specimens the existence of
series of zocecia is obscured, as the zocecia in many cases seem to be
scattered over the stem and the whole length is not usually
covered; but an examination of the stolon of these N aples speci-
mens after the zoecia have been removed, shows two groups of
about four longitudinal rows of rosette-plates, so that the differ-
ences between the Naples and Zanzibar specimens are but slight,
although the conditions of luxuriance are very different. Huincks’s
name of biserialis and } MacGillivray’s bilateralis would have been
very suitable for this form. There are frequently more than
three branches at the end of the internode, sometimes as many
as SIX.
The zoecia are about 0°3 mm. long, and in the sections made
the ova are usually single, or in some cases there are two in an
ovarium.
The embryo is surrounded by an ovicell sac, much the same as
in Adeonelle * and in Diplodidymia*, the crowth of the embryo
and of the sac going on simultaneously, and in many cases they
are so close together that at first it is difficult to distinguish the
sac from the embryo. In these cases the ovum must pass to the
distal end under the operculum, from where it is developed.
On the other hand, in most of the Cheilostomata and in the
Cyclostomata, the ovicell is formed before the embryo is ready for
it. In the Cyclostomata the walls may be seen starting at various
points f to ultimately unite to form the ovicell.
The rosette-plates were first described by Reichert? in this
species, and it does not seem that anyone, except Smitt, had
previously described anything of the kind, but what Smitt figured
* Waters, “ Bry. from Zanzibar,” Proc. Zool. Soc. 1913, p. 529, pl. Ixxili. figs. 3
& 5, and p. 490, text-fig. 79.
+ ©On the Ovicells of some Lichenopore,”’ Journ. Linn. Soc., Zool. vol. xx. pl. xv.
fig. 6 (1888); “ Mar. Biol. Sud. Red Sea, Bry..” Journ. Linn. Soc., Zool. vol. xxxi.
. xxv. fig. 16 (1909).
E: Vere. Anat. Untersuch. ti. Zoobotryon pellucidus, Ehr.,” Abhand. k. Akad. der
Wiss. Berlin, p- 276 (1869).
850 MR. A. W. WATERS ON
as a communication pore was evidently the entire plate, it not
being realised that there were not large pores, but only minute
perforations. Reichert described and figured the plate as with a
central perforation with 8-10 pores round it * (pl. iii. fig. 7), but
this is not the case, as there is only one? perforation, and what
Reichert took to be pores surrounding a central pore are really
a cirele of cells, on the older side of the rosette-plate, with a
relatively large round nucleus anda thin prolongation. The disk
is thinner than the surrounding walls, as figured by Reichert
(loc. cit. pl. iii. fig. 7). Sections and preparations of Zoobotryon
from Naples, the Soudan and Wasin, enable me to add slightly to
our knowledge of this species.
By the rosette-plates of the zocecia the cells just mentioned
often stain very darkly, so that no structure can be seen, but in
other cases the plate is seen with all these cells in a semicircle =
(PI. III. figs. 7, 8, 9, 12) pointing their thinner edge to the centre,
or with them even raised to the one minute opening. Above,
that is on the younger side of the rosette-plate, there are a
number of cells, probably usually the same number as below, anc
these also have long thread-like projections which pass to the
opening to touch the projections of the under circle of cells, that
is as if eight fingers from below were raised to touch eight fingers
from above. From both sets of cells spread other cells in a
more or less radiating manner, and to these mounds of cells the
funicular or plasma-threads reach, distinguishable from the others
by their elongated nuclei. These plasma-cords spread to all the
organs of the zoarium.
The layer of mesenchym-cells lining the zocecial walls (PI. II.
fig. 10; Pl. IV. figs. 2, 3) spread to this mound of cells, and can
be distinguished by the round cells and round nuclei from the
funicular threads with their long cells and elongate nuclei. There
is sometimes on one side a more or less semicircular cover over
the circle of radiating cells, which it is difficult to understand,
and they never occur in the early stages, and certainly they
cannot be continuous all over or there would be no connection
from the two sides.
An ovum is often seen pretty near to the rosette-plate, so that
it must move up to the distal end to be enclosed in the ovisac.
When there is an ovum, and usually in the older zoocia, there is
a considerable change in the funiculus near to the rosette-plate,
as it has become granular (PI. IIT. figs. 10, 11), so that from this
appearance the condition of the zocecium can be surmised.
Nitsche$ spoke of the accumulation of cells over the rosette-
plate as a “‘Pfropf” (plug or stopper), and Reichert mentioned
* This figure was copied by Hincks, Brit. Mar. Poly. p. ix, fig. 4.
+ Joliet, Bry. des Cotes de France, p. 31, footnote, says that Bowerbenkia has only
one perforation. AsI have previously stated, Zoobotryon and Bowerbankia should
not be generically separated.
t This is the “joneturie” of Jullien, who describes a similar arrangement of cells
to the joncturie of Schizoporella malusii Aud., ‘Cap Horn,’ p. 42.
§ ‘ Beitr. z. Kennt. der Bryozoen,” Zeit. f. wiss. Zool. vol. xxi. p. 9 (1871).
BRYOZOA FROM ZANZIBAR, 851
the thickening of the “‘Communale Bewegungsorgan ” above and
below the septa, whereas F. Miiller od ealliadl it a ganglion,
maintaining that there was a common nervous system “through
the colony; also Smitt *, in Bugula, called them colonial nervous
ganglia. We have seen that to these mounds of cells the network
of plasma threads reaches, and this 1s what has been ealled the
colonial nervous system and the colonial organ of movement, and
it was figured by Miller, Reichert, and Nitsche-as a stout, solid
or tubular body, but Henetiendl there are a number of anastomosing
and reticulate threads, though with very low powers it may look
like a band: neither alive nor in stained sections is any such
solid or tubular band seen. Nitsche’s figures must be looked
upon as diagrammatic,and Reichert’s, as I have previously stated,
as though he had Zoobotryon in a pathological condition, and it is
to be regretted that Hincks copied his figures. Vigelius, Freese,
and others have correctly understood these plasma threads.
It is strange that no histological work has been done on
Zoobotryon, as it is eminently suitable for elucidating many
interesting and important points of cell-structure, etc.
The name rosette-plate was given supposing that there was a
rosette of pores, whereas there is really a rosette of cells.
Loc. Add: Florida (Osb.); Chuaka Bay, Zanzibar, 2 fath. (509),
collected by Crossland.
BowERBANKIA PuSTULOSA (Ellis & Solander).
Sertularia pustulosa Ellis & Solander, ‘‘ Nat. Hist. of many
curious and uncommon Zoophytes,” p. 54 (17&6).
Bowerbankia pustulosa Hincks, Brit. Mar. Poly. p. 92
pl. Ixxvi. figs. 1, 5 (1880); Calvet, “ Bry. Ectoproctes,” pl. vi. fig. 13,
pl. vii. fies’ 4- 8, Ole WIM, Teva, AN Golly axe ibe, Wey yolks oa, figs. 18, is
pl. xu. figs, 15- i, jolle, sani figs. 11, 14 (1900).
The descriptions and determinations of Lowerbankia are so
uncertain that there seems little object in giving a full list of
synonyms. This species has been mistaken 7 for Valkeria uva L.
and 5. imbricata Adams.
Loc. British; Mediterranean ; Chuaka, Zanzibar Channel, shore
(523), collected by Crossland.
MIMOSELLA BIGEMINATA, Sp. n. (PI. IIT. figs. 1-3.)
The zoarium eonsists of unbranched stems (about 0°3 mm.
diameter) rising from a spreading stolon (about 0°15 mm.). The
first internode of the stem is long (say about 1:65 mm.) followed
by one about 0-4 mm., and then the remaining ones are shorter,
say 0°35 mm. There may be as many as 50 internodes in a stem.
In the lower internodes there is only a pair of opposite zocecia
close to the distal end, but after the second or third internode there
is a second pair placed rather to the side and lower down but
* Hafs-Bry. Utveck. Ofv. Stockh. Akad. xxii. pl. vi. fig. 7 (1865).
+ See Waters, “ Mar. Biol. of the Sudanese Red Sea, Bryozoa,’ Journ. Linn. Soc.,
Zool. vol. xxxi. pp. 249, 250 (1910).
852 MR, A. W. WATERS ON
close to the other pair. There are several cases where there has
apparently been an injury, and a new branch grows from the side
of the broken one, and then, although the original stem has had
the double pairs of zocecia, yet the new growth may have more
than two internodes with only a pair of zocecia, but subsequently
there are two pairs to each, though special causes may occasion
irregularities. The diaphragm at the base of a zoccium has one
pore. Near the base of the zocecium there are strong muscles for
moving the zocecium, and similar muscles occur in Jf, gracilis H..,
although Hincks said there were none. The new species differs
from WM. gracilis H. in the stems not branching, and in having
four zoeecia grouped at the distal end of the internode.
Triticella armata Verrill has the zoarial growth very similar to
that of Mimosella, but, according to Osburn*, it has a gizzard,
and evidently does not belong to the present group.
The name bigeminata was suggested by Dr Harmer, who, when
T told him that I had found aot figured this Wimosella, thought
that he had also found it in the ‘Siboga’ material. When he
showed me- his specimens, which are from better material than
mine, the identity seemed to me quite clear, and, on seeing mine,
Dr. Harmer agreed that this was the case, so that, with his
permission, the name originally proposed has been changed.
Loc. Ras Osowamembe, Zanzibar Channel, 10 fath. (504),
collected by Crossland.
There is a small fragment of another J/imosella from Chuaka,
which I hesitate to name as it is incomplete. Just below the
diaphragm of the main stem there is a lateral stem on each side,
and each of these has two or three single zoecia growing direct
from the stem and directed distally, with this free unoceupied
stem continuing beyond the zoccia. On the lateral stem there is
a diaphragm before and after each zocecium. In one lateral stem
there are two plates of attachment, as if there had been two pairs
of zoeecia, though on all the others the zocecia are uniserial. The
zovecia are about 0°25 mm.—0°3 mm. long. The stem is about
0-02 mm. diam. The growth of this species somewhat reminds us
of Farrella atlantica B., which, however, has the zocecia more or Jess
stalked, whereas in this Wimosella the base of the zocecium is
rounded like that of Bowerbankia. In Valkeria uva L. there is a
diaphragm above and below the group of zocecia, whereas in
Mimosela gracilis H., F. atlantica, and this species ‘there i is only
the one diaphragm just beyond the branches.
FarrELLA ATLANTICA Busk. (Pl. IV. fig. 9.)
Farrella atlantica Busk, ‘Challenger’ Exped., Zool. vol. xvii.
p. 37, pl. vu. fig. 3 (1886); Thornely, ‘‘ Rep. Pearl-Oyster
Fisheries of the Gulf of Manaar,” p. 128 (1905); ‘Mar. Poly.
Ind. Ocean,” Trans. Linn. Soc. vol. xv. p. 157 (1912).
* Osburn calls it Hippuraria, but I have shown that the genus was founded on a
mistake, as the “stem” was a seaweed upon which it grew: see “ Rep. Sudanese
Bry..” Journ. Linn. Soc., Zool. vol. xxxi. p. 241,
BRYOZOA FROM ZANZIBAR. 853
Hippuraria verticillata Hincks (non Heller), Ann. Mag. Nat.
Hist. ser. 5, vol. xix. p. 311, pl. ix. fig. 8 (1887).
I have only found a small piece from the Red Sea, but it 1s
evidently widely distributed, as I have it from Naples, the Red
Sea, and Zanzibar.
The rhizome of the Zanzibar specimens is 0°01 mm. in diameter ;
the zocecia are 0-4 mm. long and 9-1 mm. wide, about the size
given by Busk. At intervals on the stalk there is a pair of
zocecia growing from an expanded part, and there is often from
the same expansions a pair of radicles growing at right angles to
the stalk; also frequently there are lateral branches near to the
expansions, and these sometimes have pairs of zocecia near to the
main branch. |
A zoecium from which a polypide has disappeared usually
assumes a Swollen * barrel-shape (PI. LV. fig. 9, 6), as the muscles
which kept them in shape have now disappeared, and the same
kind of thing occurs in many Ctenostomata.
IT am unable to follow Hincks when he identifies his Hippu-
raria verticillata + with Heller's species; also I have shown that
the genus Hippuraria was founded upon a mistake, and is only a
synonym of 7'ridécella. Although Hincks speaks of a group of
cells at the joint, the figure shows a pair, and perhaps other nearly
adjacent pairs have given the appearance of a group, so that,
although I have scarcely any doubt as to Hincks’s verticillata
being a synonym of Busk’s atlantica, there is just the possibility
of this not being the case.
The peduncle is attached to the zowcium excentrically, and
there is a muscle from the side of the wall to the projecting base
of the zocecium by which it is moved as a whole.
There is a somewhat similar muscle in Mimosella gracilis
Hincks. Joliet mentions this also in his Valkeria nutans =, and
Hincks was apparently unaware of Joliet’s later description when
describing H. verticillata. Heller’s figure of Valkeria verticillata,
and Hincks’s figure of Valkeria wva in the ‘ British Marine
Polyzoa’ are very similar.
Loc. Bahia, 10-20 fath. (Chall.); Naples (4. W. W. coll.) ;
Adriatic ; Suez; Ceylon (Z7%.); Indian Ocean (7h.); Amirante,
25 fath. (7h.); Providence, 50-78 fath.(Zh.). Ras Osowamembe,
Zanzibar Channel, 10 fath. (504); Wasin, Brit. E. Africa, 10 fath.
(500), collected by Crossland.
VALKERIA UvA (Linneus).
For synonyms see Waters, ‘‘ Mar. Biol. of the Sudanese Red
Sea, Cyclostomata, Ctenostomata, and Endoprocta,” Journ. Linn.
Soc., Zool. vol. xxxi. p. 250, pl. xxiv. fig. 13, pl. xxv. figs. 4,
12, 13 (1910); Osburn, “Biol. Survey of Woods Hole and
* Journ. Linn. Soc., Zool. vol. xxxi. p. 239, pl. xxv. fig. 6.
+ Ann. Mag. Nat. Hist. ser. 5, vol. xix. p. 311, pl. ix. fig. 8 (1887).
£ “Etudes Anat. & Emb. sur le Pyrosoma giganteum,” p. 106, pl. v. fig. 4 (1888).
854 MR. A. W. WATERS ON
Vicinity,” Bull. Bureau of Fisheries, vol. xxxi. pt. 2, p. 606
(1911).
The specimens from Chuaka have groups of zocecia at distant
intervals, and there are fully formed embryos in the zocecia.
Loc. See above and add: Vineyard Sound, 6-8 fath. ete.
Chuaka, Zanzibar, 2 fath., on seaweed with Starpanie dendro-
grapta Waters (508); aud Chuaka shore, on Amathia lendigera
(521): collected by Crossland.
CYLINDR@CIUM GIGANTEUM (Busk).
Farrella gigantea Busk, Quart. Journ. Micr. Se. vol. iv. p. 93,
pl. v. figs. 1, 2 (1856).
Cylindreecium giganteum Hincks, Brit. Mar. Poly. p. 535,
pl. Ixxvii. figs. 3, 4 (1880).
The largest zoccia from Chuaka are about 0°25 mm, long.
There is no dilation at the base, and the contents of the wall are
mostly calcareous, giving it a white appearance, so that at first
I was inclined to Teal it var. album. When placed in acid it
becomes transparent. The determination of Cylindrecium is
always very unsatisfactory, and probably C. gigantewm and
C. dilatatum have not always been correctly distinguished.
The stolons branch in various directions.
Loc. British; Mediterranean; Red Sea; off Portugal; Ceylon ;
Cargados ; Farquhar Reef (7’.); Queen Charlotte Island ; Tor-
tugas. Chuaka, Zanzibar, 2 fath. (508); Prison Island, Zanzibar,
collected by Crossland.
Busk1A NITENS Alder.
Buskia nitens Alder, Q. J. Micr. Sc. vol. v. p. 24, pl. xiii. figs. 1,2
(1857); Hincks, Brit. Wars Poly. p. 532, pl. iin figs. 6, 7, Onn
cut, fig. 28 (1880) ; Ann. Mag. Nat. Hist. ser. 5, ak xili. p. Os
Levinsen, ‘“ Zool. Danica, Mosdyr,” p. 83, pl. viii. figs. 12, 13
(1894).
From Ras Osowamembe, growing on the stalk of I/imosella
bigeminata, nov. Levinsen says that there are 8 tentacles.
Loc. Arctic; Davis Strait, 100 fath. ; British; Danish ; Queen
Charlotte Island (#.). Ras Osowamembe, Zanzibar Channel,
10 fath. (504), collected by Crossland.
PEDICELLINA SPINOSA (Robertson). (Pi. IV. figs. 10, 11.)
Myosoma spinosa Robertson, “Studies in Pacific Coast Euto-
procta,” Proc. Calif. Acad. of Sciences, ser, 3, vol. il. p. 324,
pl. xvi. figs. 1-12 (1900).
There are two specimens from Wasin, which well show most of
the characters mentioned by Dr. Alice Robertson. ‘The spines
en both the stalk and calyx are stout and long, whereas those on
P.cernua may be called almost hairs, and the spines are abundant
on the one side and absent on the other, the obliquity of the ten-
tacular region also occurs. The number of tentacles is about 14.
BRYOZOA FROM ZANZIBAR. 855
The stalk is broad, especially at the base, and is also wide
where the calyx is attached, though not so wide as figured by
Robertson in many of the figures, but she shows a distinct
separation in fig. 4. In the stolon there is a septum pretty near
to the stalk on each side of it, and there is a considerable space
from stalk to stalk.
‘The preparation was stained and mounted for some other more
transparent species, and it is not suitable for following any
inuscle from the stalk up the calyx, nor am I able to see anything
suggesting it.
While unable to aceept the genus Myosoma at present, of
which I have been unable to make sections, it may turn out that
it is advisable to retain the genus.
1 now think that the Pedicellina from Naples with numerous
stout recurved spines, to which I referred*, is P. hirsuta Jull.
Loc. Tomales Bay, California, beach; Fort Point and San
Pedro, California. Wastin, Brit. E. Africa, 10 fath., with Coral-
lina etc. (500), collected by Crossland.
BARENTSIA GRACILIS (Sars).
For synonyms see Waters, ‘“‘ Rep. Mar. Biol. of the Sudanese
Red Sea, pt. 11.,” Journ. Linn. Soce., Zool. vol. xxxi. p. 251 (1910)
and add :—
Pedicellina gracilis Pottinger, ‘“‘ Anat. des Pédicellines de la Cote
d’Ostende,” Arch. de Biol. vol. vil. p. 300, etc. (1886).
Ascopodaria gracilis Norman, “'The Polyzoa of Madeira and
neigh. Isl.,” Journ. Linn. Soc., Zool. vol. xxx. p. 277 (1909).
Some of the stalks have the swelling in the middle, which have
been found from many places, and in consequence of which the
species nodosa was made. James Ritchie? has confirmed what
I have said ¢ as to the genus Gonopodaria being superfluous.
Loc. Arctic; European coast, generally; Mediterranean ; Red
Sea; Madeira; Australasia. Ras Osowamembe, Zanzibar
Channel, 10 fath. (6504); Wasin, Brit. EK. Africa, 10 fath. (500) ;
Chuaka, Zanzibar, ‘from growth on elytron of Aphroditid” ;
Chuaka, Zanzibar, 2 fath. (508), (509), (519): collected by
Crossland.
LoxosoMA SINGULARE Keferstein.
Lowxosoma singulare Kef. Zeit. wiss. Zool. vol. xii. p. 13, pl. xi.
fig. 29 (1862); and add to Miss Jelly’s Catalogue :—
Loxosoma singulare Jull. & Calvet, “ Bry. prov. de l’Hiron-
delle,” p. 28, pl. 1. fig. 5(1903) ; Harmer, ‘‘ Struct. and Devel. of
Loxosoma,” Q. J. Micr. Se. 1885, p. 4 .
There are a number of specimens growing on Schizoporella
p. 252 (1910).
+ “On an Entoproctan Polyzoon (Barentsia benedenz) new to the British Isles,”
Trans. Roy. Soc. Edin. vol. xlvii. p. 835 (1911).
t “ Résultats du Voy. du S.Y. * Belgica’—Bryozoa,” p. 100 (1904); “ Mar. Biol.
Sud. Red Sea, pt. 1,” Journ. Linn. Soc., Zool. yol. xxxi, p. 252 (1910),
856 MR. A. W. WATERS ON
nivea B.,and they are mostly, or at any rate exceeding frequently,
attached to the operculum, indicating that the movement gained
in this way is favourable to the Loxosoma.
Loc. Holland (Kef.); Shetland (Hincks); Naples (Harmer,
etc.); Newfoundland (J. & C.). Prison Island, Zanzibar Channel,
8 fath. (505), collected by Crossland.
ADDENDUM.
Since I wrote about Lagenipora socialis H. in the description
of the Cheilostomata from Zanzibar, pt. i. p. 510 (1913), I have
examined the Norman Collection, recently sent to the British
Museum, and there is a specimen from Hastings, sent by
Mr. Hincks to Canon Norman as Lagenipora socialis, which has
a pore at each corner of the ridge, as I described in the Guernsey
specimens.
This entirely confirms the view that Lagenipora socialis is the
type of the group which [I have several times maintained was
Lagenipora, but which Levinsen has called Siniopelta. By this
specimen it is now definitely settled, and does not admit of
further question, but the examination has brought out another
interesting point. The Cedleporella Norman belongs to the same
genus, which, however, Norman at some time recognised, for he
wrote on Hincks’s co-type of Lagenipora socialis, “ Celleporella
lepralioides.” However, under C. lepralioides he had two species,
first the Z. socialis, and then from Guernsey and from Hardanger
Fiord specimens with several pores by the ridge of the ovicell,
which are probably Z. lucida, as well as L. socialis, both of which
he had identified with Celleporella lepralioides. Of course, Celle-
porella has to disappear, as it was not recognisable from the
description and figure.
I might have mentioned in the same paper, when speaking of
Actea, p. 464, that what I call the bulging out of the zoecial
wall, for the ovum, before the ovicell has been formed, has been
figured by Prouho in Cylindrecium™.
In looking over seaweed for Pedicellina a few zocecia of Lepralia
poissonit Aud, have been met with from Wasin, Brit. EK. Africa,
10 fathoms. The primary zocecium has two or three more spines
than Levinsen’s figures. The species is known from the Atlantic,
Australasia, Indian Ocean, and Japan (A. W. W. coll.), and is a
common species in many localities. It also oceurs fossil.
From Chuaka, 3 fathoms (523), a few zocecia of Beania inter-
media Hincks were found. There are no frontal spines, and near
the proximal end there is a large plate for the attachment of the
radicle, and at each side of this, near the border of the zoccium
either at the same level as the radicle or higher up, there is a
* Contrib. a Vhist. des Bryoz. »” Arch. Zool. Ex ér. gme ser. vol. e A
figs. 14-17 (1892). : P x, pl. xxiv.
BRYOZOA FROM ZANZIBAR. 857
tubular projection, and from one of these a new zoccium may
grow. Close up to the distal border on the dorsal surface there
is on each side a small pore, which, however, is not seen in the
Chatham Island specimens. Now I have previously remarked
that most ‘* Beanie have six tubular connections, whereas this
has only four,” so that there is reasonable ground for considering
these as vestigial, thus representing six connections. Two pores
are found in this position in several Cheilostomata, as Brettia,
Cateicella, etc., and we may, perhaps, now see the significance of
these spots or disks.
The zocecia are about a third larger than those from the
Chatham Islands and twice as large as those from New Zealand.
The B. intermedia has been found in New Zealand, Tasmania
(H.), Chatham Islands (W.), Red Sea (W.), Australia, Indian
Ocean.
EXPLANATION OF THE PLATES.
Prate I.
Fig. 1. Crisia inflata, sp.n. 25. From Wasin.
Do. do. X 85. Showing ovicell.
. Crisia elongata M.-Ed. X12. From Wasin.
Do. do. Natural size.
Crisia sertularoides Aud. X25. From Wasin.
Do. do. x 3.
Crisia circinata, sp.n. X12. From Ras Osowamembe.
Do. do. X about 2.
Do. do. X 25. Ovicell seen from the side.
G2) C9) SSH Gm ONE COS)
Puate If.
Fig. 1. Entalophora wasinensis, nom.n. X 25. Showing ovicell. From Wasin.
Do. do. x 60. Section of ovicells containing embryos.
Do. do. X 330. Embryo.
Do. do. X 85. Section of ovicell containing embryos.
. Idmonea interjuncta MacG. X12. Showing ovicell with oceciostome (oe.).
From Ras Osowamembe.
. Idmonea radians, var. erecta Busk. X25. Showing ovicells with ocecio-
stome (oe.). From Ras Osowamembe.
Do. do. do. X 25. Longitudinal section of the ovieell. Cut
parallel to the anterior and dorsal surfaces.
Do. do. do. Natural size, showing the position of the ovicells
Fig. a. Idmonea radians Lam. typica. Natural size.
. Entalophora wasinensis, nom.n. X 330. Section of the surface pore-tubes ;
(m.) exterior membrane.
. Idmonea radians, var. erecta Busk. X85. Section from the anterior to
the dorsal surface of the zoarium, showing the lobes of the ovicell and
the ooeciostome (0e.) as well as the polypides (p.). The structure of the
wall of the ovicell is seen at the left (s¢.) with one or two rosette-plates
at the base of the broad pore-tube. There is an outer membrane (m.).
Oo DM ST D MBO
a
i=)
Puate III.
Fig. 1. Mimosella bigeminata, sp.n. X85. Showing the lower internode with
only a pair of zocecia, while the upper one has two
pairs. From Ras Osowamembe.
2. Do. do. X 6. Showing several stems growing from the creeping
stolon.
3. Do. do. < 85. Showing a new stem growing from the side of
an old one after mutilation,
858 ON BRYOZOA FROM ZANZIBAR.
Fig. 4. Zoobotryon pellucidum Ehr. 250. Embryo in ovisac (0s.). Zocecial
wall (zw.). This shows the embryo hollow, which
is very generally the case in embryos of this type.
5. Do. do. < 200. Disk of rosette-plate showing one small pore
in the centre.
6. Do. do. < 375. Disk of rosette-plate with a circle of cells
round the pore.
Ac Do. do. X 700. Section across the rosette-plate showing cells
directed to the small opening; with meseuchym
cells above. Only two of the radiating cells above
the rosette-p'ate are shown, as the others are only
seen in a different focus, but there will be 8 or 9.
8. Do. do. X 330. Section across the rosette-plate in a septum of
amain stolon. Showing a spreading plasma with
cells scattered about. From the Sudan.
9. Do. do. x 700. Section across the rosette-plate from the stolon
to the zocwecium.
10. Do. do. X 330. Section across the rosette-plate; above it there
is an irregular granular mass. ‘This is in a zoceciuim
in which there is an ovarium, and degeneration has
taken place.
iit Do. do. X 330. Section across the rosette-plate from a stolon
to a zocecium, showing irregular granular masses in
a zoccium that is degenerating.
12. Do. do. < 330. Section across the rosette-plate from a stolon
to a zocecium.
In all cases the figures show the older part below the rosette-plate, and the
younger above.
Puate LV.
Fig. 1. Amathia vidovici Hell. X 12. From Wasin.
2. Do. do. >< 330. Section where a new branch is formed on each
side, showing the cells round the two rosette-plates.
The plug of cells on the lett are not cut close to the
opening of the rosette-plate, and the two inner plugs
have become somewhat granular.
3. Amathia lendigeraL. 500. Section through the rosette-plate of the
septum of the stolon. Examined with 1/12
immersion. From Swanage.
4. Do. do. X 25. Showing the branch at a bifurcation arising
almost at right angles to the main stem.
5. Crisia denticulata Lamk. X 330. Section through the inner wall of a
zocecium, showing the connections from zocecium to zocecium (¢.). From
Swanage.
6. Crisia elongata M.-Ed. X 250. Proximal end of zowecium—(a.) showing
connection to the two neighbouring zoccia, through numerous tubes in
which there is a septum in the middle, zocecial walls (w.). From Wasin.
7. Stomatopora. Vrimary zowcium. X 85.
8. Tervia irregularis Meneghini. Comb-like process in the zoccial tube near
where it becomes erect. From Naples.
9. Farrella atlantica Busk. X85. From Ras Osowamembe.
10. Pedicellina spinosa Robertson. X 85. From Wasin.
11. Do. do. x 25.
12. Zoobotryon pellucidum Ehr. Natural size. From Chuaka Bay.
ON AVIAN CESTODES. 859
47. Contributions to the Anatomy and Systematic Arrange-
ment of the Cestoidea. By Frank H. Brepparp, M.A.,
D.Sc., F.R.S., F.Z.8., Prosector to the Society.
[Received May 65,1914: Read June 9, 1914. ]
(Text-figures 1-11.)
XIV. ON A NEW SPECIES OF RHABDOMETRA, AND ON THE
PARUTERINE ORGAN IN O7IDIT_-ENIA.
INDEX.
Page
Description of Rhabdometra cylindrica, sp. n. ............... 859
Paruterine Organ of Otiditenia ewpodotidis Beddard ...... 879
I have recently obtained a considerable number of examples of
a Tapeworm from an African Partridge, Caccabis melanocephala,
which [ refer, temporarily at least, to the genus Rhabdometra,
though a closer comparison of this worm with the known species
of Lhabdometra may ultimately necessitate its separation from that
genus. The worm occurred in the Partridge associated with
several other species, and I found always a relationship in numbers
between the different forms which inhabited that bird. I examined
altogether five specimens of the Caccabis in three of which were
found examples of the Rhabdometra. In two examples there were
a large number of specimens of the Rhabdometra, and in the other
example only two specimens. The birds, which were infested by
many specimens of Ahabdometra, contained also apparently only
one representative of other species of Tapeworms. In the one
ease it was a Davainea and in the other a Cotugnia (¢?) In the
bird where there were only two Lhabdometra there were many
Davainee.
The general appearance of this worm is indicated in text-fig. 1.
It is long and slender; when alive the longest examples were
fully six inches or so in length. In spirit the dimensions are
somewhat lessened. This great length is accompanied by only a
small diameter; the width varies from something under one
millimetre, to a little over a millimetre. The greatest diameter
was anteriorly in the body but some way behind the head; here
the proglottids are broader than long. ‘The scolex as a rule
appears to be a little less in diameter than the ensuing region of
the .strobila. But occasionally, as depicted in text-fig. 1 A, the
scolex stands out as a globular body followed by a constriction.
In an examination of the living worms I noted one example in
which there was no neck, an unsegmented region following upon
the scolex ; and I have one mounted preparation (represented in
text-fig. 2), where the same feature is to be seen. But I have no
note as to whether these examples were the same. As a general
Proc. Zoou. Soc.—i914, No. LVIII. 58
860 DR. F. E. BEDDARD ON
rule there is no neck and segmentation begins immediately after
the scolex. The scolex is rather small and the suckers are large
in proportion, occupying most of the area of the scolex. There
was no trace of any hooks whether related to a rostellum or upon
the suckers. I ascertained this by transverse and longitudinal
sections as well as by the examination of entire scolices mounted
Text-figure 1.
A. Head-end of an example of Rhabdometra cylindrica.
B. Another and complete specimen of the same species.
in glycerine. It is very important to be positive upon this point,
since the difference between the genera habdometra and Paru-
terina is practically entirely to be found in the absence in one and
the presence in the other of a rostellum armed with hooks. The
strobila consists of proglottids, which are at first narrow and then
AVIAN GESTODES. 861
rather wider, ‘The end of the body, often more than one half of
the entire worm, is made up of very long proglottids which I
found to measure as much as five and six millimetres in length.
They are thus quite as much as or even more than six times as
long as they are broad. There is no one of the known species
of Rhabdometra in which the ripe proglottids are so long as in
the species described in the present communication. It may be,
Text-figure 2.
Anterior end of two specimens of Rhabdometra cylindrica.
The vight-hand figure shows the existence of an unsegmented neck rare
in the species.
however, that the individuals of the various species examined by
my predecessors were not so fully mature as those which I have
myself been able to study. The greatest length of the hinder
proglottids is to be found, as it would appear, in Lhabdometra
nigropunctata, where, according to Crety*, they are three times
the length of their diameter. That is considerably less than I have
* Boll. Mus. Torino, v. 1890, No. 88. It should also be noted that this species,
which measures 140 mm., is nearest in length to mine.
58*
862 DR. F. E. BEDDARD ON
met with. The anterior relatively broader segments are rather
flattened in transverse section. The posterior proglottids are
sometimes quite cylindrical in form, being circular in section. Or
they may be more oval, though still of great depth dorso-
ventrally.
In transverse sections such as are represented in text-fig. 6,
the cortical layer is seen to be fully as thick as, and occasionally
even a shade thicker than, the medulla. The distribution of the
longitudinal muscular layer seems to be very much as it has been
described by Fuhrmann* for the allied species i. numida. Next
to the transverse muscles there is a layer of rather widely spaced
bundles with not more and sometimes less than 5 or 6 fibres in
each. Between these and the subcuticular layer are numerous
scattered longitudinal fibres implanted singly. The transverse
fibres form a rather thick layer, and in the anterior region of the
proglottid, where the medulla is free from the testes ete., and in
the area partially occupied by the paruterine organ, the trans-
verse fibres encroach further upon the medullary parenchyma,
which is largely pervaded by them. Dorso-ventral fibres are also
abundant in the medullary layer and, as mentioned later, are not
at first displaced by the paruterine organ.
The water-vascular system consists of the two usual longi-
tudinal vessels, of which the very much larger ventral vessels are
united by an equally wide transverse vessel at the posterior end
of each proglottid. The opening of the longitudinal vessel into
the transverse vessel is guarded by a valve which prevents the
reflux of fluid into the longitudinal vessel anteriorly. I could find
no other branches of the ventral vessel to form a network such as
appears to exist according to Ransom in habdometra nullicollis.
The dorsal vessel lies above the ventral and often rather to the
inside ; it is very small but has relatively thick walls. The
genital ducts pass between the two vessels and below the nerve-
eord, which is displaced towards the dorsal side from its usual
position, where the ducts pass beneath it. It will be noted that
in the relations of the nerve-cord to the genital ducts the present
species differs from both kh. nudlicollis and Kh. similis, where
the genital ducts pass dorsal of the nerve-cord.
The testes are numerous, and often very closely pressed to-
gether. ‘They lie on all sides of the ovary and vitelline gland,
being found laterally and posteriorly as well as anteriorly in the
segment. In the most mature segments the testes are only plainly
to be recognised posteriorly in the segment lying behind the
uterus. Laterally the testes in the ripe but not fully mature
_proglottids extend as far as the water-vessels on each side. An-
teriorly the boundary of the area occupied by the testes is some
way short of the anterior margin of the proglottid. In transverse
sections of proglottids where the uterus is nearly fully developed,
the testes are seen to lie dorsally of the uterus, which latter organ
* Res. Swed. Zool. Exp. Egypt, pt. iii. No. 27, 1909.
AVIAN CESTODES. 863
occupies the whole ventral surface of the proglottid. The ex-
tension of the testes anteriorly in the segment seems to be
prevented by the paruterine organ, which in those proglottids
where the testes are fully ripe extends backwards for about half
the length of the proglottid and takes up the greater part of the
available space. It is important to insist upon the fact that
the testes surround the female gonads, since this does not appear
to be the case with other species of this genus Rhabdometra as
figured by Ransom. The genus is, in fact, partly defined by the
existence of the testes only behind and at the sides of the female
organs. The testes are only two deep in a given segment.
The cirrus-sac is long and slender, longer than that of the
species figured by Ransom, but considerably shorter than the
cirrus-sac of RA. nwmida of Fuhrmann, which reaches to the middle
of the segment. In segments where the gonads are ripe but in
which there is as yet no uterus, the cirrus-sac very nearly reaches
the middle of the segment; but in older and wider segments it
only just crosses the ventral water-vessel. Its course is obliquely
forward from the point of opening on to the exterior, which is
vather behind the middle line of the proglottid. There can
hardly be said to be a genital cloaca : a funnel-shaped depression
of the body, into the bottom of which the genital ducts open (the
male duct anterior to the female), not showing the characters of
a distinct chamber such as that of, for example, Hugonodeum.
In horizontal sections the cirrus-sac often has a serpentiform out-
line, being like an elongated S$. Occasionally it is slightly dilated
at the internal end, but the cirrus-sac of this Rhabdometra never
has the bottle-like form of that of many other tapeworms. In
consequence of its length and slenderness the cirrus is not much
coiled within it, lying mostly straight or rather in an undulating
line. I could detect no spines upon the cirrus when protruded.
I did not observe any autocopulation. The walls of the cirrus-
sac are thick and very muscular. The internal coat is of cir-
cularly running fibres, the outer layer fibres run longitudinally.
From the internal end of the cirrus a retractor muscle runs for
some way into the medullary tissue. This character is also found
in other species of the genus Rhabdometra. The vas deferens
forms a large coil which is situated posteriorly to the paruterine
organ, and extends backward in the pvoglottid to as far as the
receptaculum seminis. At about this point the efferent tubules
from the testes meet it. There is no vesicula seminalis.
The ovary lies at about the middle of the segment. It is in
front of and larger than the vitelline gland:
The vagina is long and thick-walled, and outside of the mus-
cular walls is a layer of stalked glands which are deeply stained
by hematoxylin. The course of the vagina is directed parallel to
and slightly away from the cirrus-sac to begin with; it then
curves more backwards to open into the receptaculum seminis.
Its course is apt to be rather undulating. Ransom has figured a
sudden change in the character of the vagina in the species
864 DR. F. E. BEDDARD ON
described by himself, occurring at some distance from the open-
ing into the receptaculum. It here becomes much narrower.
Nothing of the kind oceurs in Rhabdometra cylindrica, except,
perhaps, just at the orifice ; and, moreover, the coating of gland-
cells extends over the entire vagina, up to the receptaculum.
The latter is large and spherical to rather oval in form; it lies
obliquely to the longitudinal axis of the body towards the pore
side.
§ The Paruterine Organ.
This structure is of such importance in the group of tapeworms
of which the present species is a member, that it needs a detailed
treatment for comparative purposes. In the living worm the
paruterine organ is exceedingly conspicuous as a rod-like body
at the anterior end of the proglottid, often of a brilliant white,
thus contrasting with the more pellucid tissues of the outer layers
of the worm. This aspect led me at first to regard the paruterine
organ as the uterus crammed with eggs, which might be expected
to show a bright white owing to the innumerable separate and
minute embryos. In the proglottids the increasing length of
the paruterine organ could readily be observed owing to its ex-
treme conspicuousness. It was so distinct from the rest of
the proglottid in its neighbourhood, that each paruterine organ
suggested a conical peg attaching two consecutive segments. In
the living worms the paruterine organ is a perfectly rod-like
structure, without any obvious twist of any kind. It was seen to
diminish slightly in width at its terminal end, and was never
seen to extend to the posterior end, though the organ appeared to
commence at the very beginning of the proglottid. The whole
appearance of the organ as seen with a lens, suggests that it is
produced by a growth from before backwards, and not vice versa.
In alcohol-preseryed examples the paruterine organ is no longer
visible in the intact worm, nor is it in specimens examined
whole after clearing but without staining. This seems to suggest
that the bright white appearance of the paruterine organ in
the living worm is caused by air-spaces in the spongy tissue of
which the organ is composed. But it must be admitted that this
of itself is difficult to understand. Still, the organ certainly has
the appearance during life of being composed of a fine froth.
I have examined the organ in the preserved worms by means of
transverse and longitudinal sections. I have already spoken of
the organ when fully developed as being rod-like, or perhaps
rather style-like, as it diminishes to one end. But in trans-
verse sections it is seen that the paruterine organ is only
rod-like, and thus circular in section, in the fully mature pro-
glottids. These proglottids are themselves tubular and oval,
or even quite circular in transverse section. In more anterior
segments the form of the proglottid is more flattened, and the
paruterine organ shares in this alteration of form. In such
proglottids the organ is more flattened and tends to have the
AVIAN CESTODES. 865
shape, in transverse sections, of a parallelogram with rounded
angles. The most important point in the development of this
organ is that it is wholly unconnected with the uterus. The
paruterine organ is found in proglottids where there is as yet no
trace of a uter us.
There can, therefore, be no comparison with the paruterine organ
of such a form as Avitellina, ‘‘ where the uterine wall cells... .
supply the origin of the egg-pouches or paruterine organ” *, With
reference to species of Ahabdometra, the statements of Ransom
are not definite. In the case of Rh. nullicollis, that author writes t
that ‘the parenchyma in front of the uterus becomes dense and
fibrous and develops into a prominent paruterine organ, which
behind is in immediate relation with the anterior end of the
uterus.” Of the paruterine organ of 7h. similis = he writes no
more positively. We may infer from Mr. Ransom’s descriptions
that the paruterine organ does not appear before the uterus and
that it may be an outgrowth of its anterior wall. In this case
there is an important difference from the species described in the
present paper, and in any case there is a difference in time of
appearance.
The earliest appearance of the paruterine organ under a high
magnification is shown in text-fig. 3. It consists mainly in an
apparent multiplication of the nuclei of the medullary parenchyma.
In any case they are more closely aggregated for a short region
in the middle of the anterior half of the proglottid. This dense
mass of nuclei—that is dense comparatively speaking—reaches
forward to the anterior border of the proglottid. But it must be
borne in mind that the actual delimitation of successive proglottids
cannot be fixed unless the wall of the transverse water-vascular
tube fixes it. In this case the paruterine organ does not reach
the anterior limit of the proglottid in which it les. In more
mature proglottids, however, the anterior margin of the par-
uterine organ is so straight a line that one cannot help thinking
that this may be the anterior margin of the proglottid, im which
case the posterior wall of the transverse water-vessel lies within
the segment in front.
I have no evidence whether the great multiplication of the
numbers of the nuclei to form the beginnings of the paruterine
organ is due to an actual multiplication, or to a crowding together
by simultaneous migration inwards from other quarters. The
nuclei of the future paruterine body show no difference from the
surrounding nuclei of the medullary parenchyma. I have said
that the multiplication of the nucleiis the main feature of the
paruterine organ on its first appearance. The only other
difference from the surrounding parenchyma is a slight opacity,
which is, I am convinced, simply due to the crowding of the
nuclei. The network structure of the medullary parenchyma is
not at first at all altered in the future meme organ. The
* Gough, Q. J. Mier. Sci. lvi. p. 375, 1911. { Loe. cit. p. 34.
+ Bull. U.S. Nat. Mus. No. 69, p. 29, 1909.
866 DR. F. E. BEDDARD ON
area occupied by the growing paruterine organ at this stage is
rather more than a quarter and Jess than a third of the length of
the proglottid. It is interesting to note that these small
dimensions apply to the completely formed paruterine organ of
Rhabdometra nullicollis *.
Text-figure 3.
ya
Portion of an anterior sezment of Rhabdometra cylindrica in horizontal
section, to show origin of paruterime body.
7. Longitudinal muscles. par. Paruterine organ. ¢. Testes. ¢.v. Transverse
water-vascular vessel.
Text-fig. 4 illustrates a portion of a transverse section of a
proglottid with a paruterine organ at a more advanced stage of
development than that represented in the last figure. The
paruterine organ is distinctly marked off from the surrounding
* Ransom, loc. cit. p. 30, fig. 22.
AVIAN CESTODES. 867
medullary parenchyma in the middle of which it lies. But
although it is definitely marked off, it has not an outer layer of
circular muscles, such as will be described presently in the
completely adult paruterine organ. ‘The principal distinction
which the paruterine organ shows in comparison with the
Text-figure 4.
Portion of a transverse section of a young proglottid ot Rhabdometra cylindrica.
The greater part of the medullary region is occupied by the paruterine organ.
e Calcareous corpuscles in the paruterine organ. d.v. Dorso-ventral
muscular fibres. ¢. Transverse muscular fibres.
surrounding medullary region is the much denser character of the
parenchyma, which no longer presents the appearance of a delicate
network with clearer circular or oval interspaces. The network
is here and there quite visible in parts, the whole tissue being much
868 DR. F. E. BEDDARD ON
more deeply stained. I take this to be due to a solidification of
the tissue by the disposition of matter in the interstices of the
orginally existing network, which is more responsive to staining
by logwood. The nuclei are more abundant than in the sur-
rounding medullary parenchyma. But they are in the same way
Text-figure 5.
Horizontal section through not fully ripe proglottid of Rhabdometra cylindrica.
par. Paruterine organ. wé. Uterus appearing as numerous partly detached
cavities. v.d. Vas deferens. w.v. Transverse water-vascular vessel.
of two kinds—larger and clearer nuclei which appear to be
myoblasts, and smaller nuclei which belong to the connective-
tissue network. A peculiar feature m the structure of the
paruterine organ at this stage, and one which is a further proof
of the view that it is a modified region of the medullary
AVIAN CESLODES. 869
parenchyma and not an outgrowth of the generative system, is
the existence of dorso-ventral muscle-fibres: these are not
elements belonging to the paruterine organ and restricted to it,
but, as is shown in the text-figure referred to, they arise outside of
and perforate it. They pass into the paruterine organ at exactly
right angles to the transverse diameter of the proglottid. A
final characteristic of the paruterine organ at this stage is the
Text-figure 6.
Ey
avvtWWicchiavee
awe Be
Cy ; 4 le oN
a
eS
%
i
fy, :
; Pym ey aii x
Wile
Transverse section through fully mature proglottid of Rhabdometra cylindrica.
par. Paruterine organ. w.v. Ventral water-vascular vessel.
larger number of calcareous bodies which lie within it and which
are most numerous where it abuts upon the uterus posteriorly.
Ransom has remarked upon the same feature in the paruterine
organ of Sphyroncotenia. The shape of the paruterine organ at
this stage as seen in horizontal sections is shown in text-fig. 5.
In the longest, and therefore presumably ripest, proglottids the
paruterine organ differs in some few particulars from its younger
870 DR. F. E. BEDDARD ON
stages. In transverse sections, as is shown in text-fig. 6, the
organ appears to be quite circular, and it lies exactly in the middle
of the proglottid. It has sometimes a perfectly conical form
tapering towards and at the end which touches, indeed protrudes
Text-figure 7.
Sagittal section through fully mature proglottid of Rhabdometra cylindrica.
par. Paruterine organ. ¢. Testes. wt. Uterus. w.v. Transverse
water-vascular vessel.
AVIAN CESTODES. “871
into, the uterus. In other cases the diameter of the paruterine
organ fluctuates from point to point, thus showing a less regular
form such as is depicted in longitudinal section in text-fig. 7.
This difference of form is probably to be explained by uneven
contraction of the worm’s body or the muscular wall of the
paruterine organ during preservation. The paruterine organ is
long, but not quite so long as the uterus at which it ends. It
therefore occupies rather less than half of the length of the
proglottid. Dorso-ventrally the paruterine organ touches the
limits of the medullary region of the proglottid; but laterally it
does not fill up that space entirely, leaving some of the original
medullary parenchyma visible right and left. There is no
question of the inclusion of any organs in the paruterine organ
such as I describe later* in what appears to be the equivalent of
a paruterine organ in the Davaineid genus Otiditenia. The
completely formed paruterine organ of this species of Rhabdometra
is sharply marked off from the surrounding tissues by a layer of
muscles disposed in circular fashion.
The existence of such an outer muscular wall to the paruterine
organ has been noted by other observers. I believe this layer to
be adventitious and for the following reasons. In the younger
stage just described there is no muscular wall at all; but the
innermost of the transverse musculay fibres tend to follow the
outline of the oval paruterine organ, though they hardly can be
said to adhere toit. The contraction, or at least the alteration, of
the form of the paruterine organ into a circle in transverse section
would tend to further a close relationship between itself and the
immediately surrounding musculature. In any case such a
relationship exists. The shape in section and the general form
of the paruterine organ together with its muscular coat are not,
however, the only points in which the adult organ differs from
the less perfected stages. The tissue which fills it is apt to have
a concentric lamellar arrangement shown in text-fig. 8; this
is also visible in sagittal sections but is not shown in text-fig. 7,
since the latter is not of a sufficiently highly magnified preparation.
It is shown, however, in text-fig. 6.
In the younger proglottids (text-fig. 5) the paruterine organ
ends up in close contact with the uterus, as has been already
mentioned. It ends, however, in a definite border which is a
straight line. In the completely developed paruterine organ
there appears to be an absolute continuity, and the connective-
tissue core of the paruterine organ melts away, as it were, in the
cavity of the uterus. It appeared to me that the caleareous
corpuscles, which are apt to be specially abundant on the paru-
terine organ at its distal end though found throughout it, are
both smaller in many cases and generally less abundant in the
older paruterine organ, It looks as if they were used up perhaps
by the growing embryos. Furthermore, the “ perforating ” dorso-
* P. 880.
872 DR. F. E. BEDDARD ON
ventral muscles, to which I have referred inthe younger paruterine
organ, are in places, but by no means always, visible in the adult,
as may be seen by a comparison of the figures given. They are
perhaps broken by the swelling of the organ to a circular form in
section, for I have seen short fibres imbedded in the connective-
tissue core.
Text-figure 8.
Oo Vv.
>
eg So) ,e
Ss
er
a |
i= re]
$e See
oe
Cd ees
@
co.
More highly magnified view of a portion of the paruterine organ in sagittal
section.
co. Tissue of paruterine organ with calcareous bodies. ov. Ripe eggs contained
in a space within the paruterine organ. ¢7. Transverse muscles.
In describing the paruterine organ of Chapmania tapika, Prof.
Fuhrmann* remarks that in that and all*forms with a paruterine
organ the ripe eggs do not pass into the paruterine organ until
the proglottids are detached and thus ready to leave the body.
Mr. Ransom? particularly remarks that in his examples of the
genus Sphyroncotenia the mature segments showed no eggs
within the paruterine organ and that the mode of their trans-
ference was thus unknown to him. This state of affairs is nearly
true also of the Rhabdometra which forms the subject of the
present communication, but not quite. In one ripe proglottid
among many which I studied, I found embryos within the
* Res. Swed. Zool. Exp. Egypt, pt. iti. No. 27, p. 19, 1909.
+ Proc. U.S. Nat. Mus. vol xl. p. 637, 1911.
AVIAN CESTODES. 873
paruterine organ, thus incidentally proving that the organ is a
paruterine organ, if any doubt could be supposed to attach to
that identification of it. The embryos were not directly imbedded
in the lax tissue of the paruterine organ but were accompanied
by a cavity within which they lay. Whether this means that
their assumption into the paruterine organ resulted in the for-
mation of a cavity by stretching or other means, or whether a
portion of the uterus was detached as a whole and engulfed by
the paruterine organ, I do not know.
$ The Uterus.
Although the appearance of the paruterine organ before the
uterus is developed proves that the former is not a product of the
latter, it does not follow that there is no connection at all between
these two organs in their origin. I believe that I have been able
to establish a connection between the two, and that the tissue of
the paruterine organ gives rise to the uterus. I am not sure
that 1 have detected the uterus in the very first segment in
which it is developed, but if not, | am not more than three or
possibly four segments out. As is often the case with tapeworms,
the uterus appears rather suddenly and in an advanced stage of
development, at least speaking relatively. Im the segment in
question, which is the second or third with a uterus, the testes
are still active and the ovary and vitelline glands in full maturity
and not beginning to degenerate. The paruterine organ is
considerably younger than that represented in text-fig. 5; in the
sections (which were nearly accurately horizontal) the outline of
the paruterine organ was nearly square, the breadth being a little
greater than the length. The uterus consists of an irregular tube
running rather obliquely across the long axis of the body. The
tube branches somewhat and there are indications of anastomoses ;
but the retiform stage of the uterus has hardly been reached.
The uterus does not extend, as it does later, in front of the
transversely running coil of the vas deferens. It does not.
therefore, come into contact with the broad posterior end of the
paruterine organ as it does in later stages (cf. text-fig. 5). There
is, however, a connection between the two which is more than
mere juxtaposition and is, therefore, of an interesting nature.
A process extends backwards from that posterior corner of the
paruterine organ which is furthest away from the pore side of the
segment. It consists, like the paruterine organ at this stage, of
little more than an agglomeration of nuclei. This process
extends backwards until it reaches the uterus with whose walls
it is continuous.
In an earlier stage, about three segments in front of that just
described, a uterus can hardly be said to exist. I detected,
however, a thread of paruterine tissue extending towards the
same side of the body ; this came into close relations with one or
two spaces containing ripe ova situated in front of the uterus,
874 DR. F. E. BEDDARD ON
and perhaps to be looked upon as the first appearance of a uterus.
I figure also (text-fig. 9) from a segment which is the one in
Text-figure 9.
Upper figures represent two horizontal sections through immature proglottids
of Rhabdometra cylindrica.
Lower figure a more highly magnified view of the connection between the
paruterine organ (par.) and the uterus (wt.).
a. Process of paruterine organ connecting it with uterus. d. Dorsal vessel.
w.v. Transverse water-vascular vessel. ¢. Testes.
AVIAN CESTODES. 875
front of that first of all treated of in the present account of the
development of the uterus of Rhabdometra. Here the connection
of the paruterine organ with the uterus happens to be plainly
visible in one seouion. and, therefore, to be more striking and
less liable to doubt than when it has to be followed out from
section to section. Furthermore, in subsequent sections, where
the uterus is more advanced and lies also in front of the vas
deferens, the tissue of the paruterine organ is seen to pass
continuously into that of the uterus, and the nuclei of the walls
of both appear to be identical.
These facts—that is to say if it be agreed that they prove a
connection between the paruterine organ and the uterus—enable
us to get over certain morphological difficulties relating to the
homologies of the uterus and paruterine organs of some other
Tapeworms.
In describing the structure of Znermicapsifer capensis* I had
to refer to an important difference in the uterus of this form as
compared with other species referred by v. Janickif to that
genus (his own). Briefly put, the difference is this: in J. capensis
there was no continuous uterus, but only a series of detached
cavities which appeared to be formed independently in the
medullary parenchyma. These cavities were formed subsequently
to the extrusion of the ova from the ovary and their scattering
through the parenchyma of the medulla. Furthermore, there
was also to be observed, and again unconnected with the ova at
first, a condensation of the medullary nuclei to form a kind of
network pervading the medulla. This network was often to be
observed in re‘ation to the ova £.
Out of this dense tissue, which ultimately surrounds the ova, is
formed the series of paruterine organs which characterise this, as
well as a few other genera of tapeworms (Davainea, Thysanotenia).
T held that the network of parenchymal tissue, out of which the
paruterine organs were formed, and the cavities in which lay the
egos singly or in groups, were not the equivalents of the branched
uterus described by v. Janicki in an allied form, /nermicapsifer
hyracis (which I removed to the genus Zschokkeella), because, if it
were, it would be a subsequent stage due to the obliteration of the
pre-existing cavity; and as the ova appeared in it later it could not
be a subsequent stage. I believe that the matter becomes clear
through the-observations which I have recorded in the present
paper. . We have in RLhabdometra, as in Inermicapsifer, a conden-
sation of nuclei to form structures or cavities to contain the eggs.
In Rhabdometra there is one extensive condensation of the kind
to form the paruterine organ and a delicate strand which extends
through part of the rest of the medullary parenchyma and would
appear to be the seat of the formation of the uterus. In my
species of Znermicapsifer there is the same condensation of the
* P.Z.S. 1912, p. 588, etc.
+ Jen. Denkschr. xvi. 1910.
t Beddard, loc. cit. text-fig. 67, p. 583.
Proc. Zoou. Soc.—1914, No. LIX. 09
876 DR. F. E. BEDDARD ON
medullary parenchyma, but it does not become much hollowed
out to form a uterus or specially condensed in one part to form a
single paruterine organ.
On the contrary, it is condensed here and there to form several
paruterine organs. In Inermicapsifer hyracis of v. Janicki, a
hollowing out of the strands of condensed tissue occurs before the
further condensation of the medullary tissue to form the numerous
paruterine organs. I am of opinion that all these rather various
Text-figure 10.
Transverse section through mature proglottid of Rhabdometra cylindrica.
ut. Uterus. w.v. Lateral water-vascular tubes.
eases are upon one common ground plan, which does away with
the at first apparent differences of importance between such
closely allied forms as Jnermicapsifer capensis and Zschokkeella
hyracis; and also relegates to the same category other forms in
which there is buta single paruterine organ. If these suggestions
thus briefly sketched out be not accepted, there still remain the
AVIAN CESTODES. 877
facts which I have described above in connection with the
formation of the uterus in Lhabdometra cylindrica. I have
mentioned that the uterus soon after its formation is distinctly
retiform, the network being irregular but very plainly to be seen
in horizontal sections. In transverse sections, the uterus is
seen to lie ventrally and to consist of variously sized cavities
detached from each other, which is, of course, the expression of
the horizontal network when seen in that aspect. In more
mature proglottids the retiform condition is not so clear and the
uterus occupies much more of the proglottid. The arms of the
network seem to have coalesced, forming an irregularly shaped
body with partial septa dividing its interior into partly detached
cavities, and with outgrowths producing a general irregularity of
surface. In the completely mature proglottids the appearance
of the uterus has again changed owing to further development.
Text-fig. 10 shows a transverse section of such a proglottid, and
the contained uterus is seen to be circular in section and thus to be
rod-like in form. Posteriorly the uterus is divided by a median
septum into two equisized halves, but further forward the uterus
is obviously single, but the interior is divided by many ingrowing
strands of the medullary tissue into a series of chambers each of
which is seen to lodge butasingle embryo. The posterior division
of the uterus into two reminds us of the uterus of Rhabdometra
mgropunctata as figured by Crety* ; it is possible, however, that the
latter form is rather to be referred to the genus Metroliasthes, in
which the uterus is an entirely double structure. The series of
changes in the appearance of the uterus in this species of
Rhabdometra is very remarkable, and seems to contrast with what
has been observed in the other species of the genus. It is
further to be noted that we have here, as a temporary phase, a
form of uterus which is now characteristic of one genus of
Tapeworms and now of another.
From the above account we can abstract the following de-
seription of the species :—
Rhabdometra cylindrica, sp. n.
Length 150 mm.; greatest diameter 1:2 mm. Scolew of less
diameter than strobila following, suckers proportionately very large ;
rostellum and hooks absent. Neck absent; posterior segments
elongated, five times their diameter, cylindrical in form. Genital
apertures alternate regularly, behind middle of proglottid. Genital
ducts pass between water-vessels and ventral of nerve-cord. Cortical
layer thick ; longitudinal muscles not in strong bundles ; mostly
implanted singly, but a row neat to transverse layer of widely
spaced bundles with from two to five or six fibres. Circular layer
* Boll. Mus. Torino, 1890. In Crety’s figure the two posterior and globular
diverticula of an anterior curved portion are alone represented as containing eggs:
I assume, therefore, that the anterior part is all of it to be regarded as a paruterine
organ,
59*
878 DR. F. E. BEDDARD ON
well marked and also occupying, in the form of scattered fibres, most
of the medullary layer in the region of the paruterine organ. Dorsal
water-vessel very small, lying dorsal of large ventral vessel, which
communicates with its fellow posteriorly in each segment by a single
transverse vessel; no excretory network. Testes surround ovary and
vitelline gland. Cirrus-sae with a strong muscular wall, and with
special retractor muscles posteriorly, long and narrow, extending a
little beyond water-vascular tubes; cirrus without spines ; vas
deferens with a large and close coil but without vesicula senvinalis.
Cirrus-sac opens into a cloaca genitalis. Vagina opens behind
cirrus-sac, terminal region wide and muscular. Uterus at first
retiform, then irregular in shape with outgrowths, later tubular
and divided into two chambers by a median septum posteriorly :
the embryos also separated by ingrowths of walls of uterus.
Paruterine organ long and styliform, commencing at anterior end
of proglottid, nearly circular in transverse section when mature,
appears before uterus.
Hab. Caccabis melanocephala.
§ Systematic Position.
It is clear that a member of the Teenioidea with a totally
unarmed scolex, with one set of genitalia in each proglottid, and
with a single paruterine organ, can only belong to one of the
three genera Rhabdometra, Anonchotenia, or Metroliasthes, or to
a new genus allied to them.
As it is, these three genera are very near together ; the generic
differences as set forth by Ransom mainly concern the position
of the genital ducts with reference to the water-vessels and the
form of the uterus. In both of these particulars the present
species is like Rhabdometra ; sufficiently so also, in other features,
to warrant its inclusion in that genus. ‘There are, however,
differences which prevent the reference of my species to any of
those which have been already described. In none of the
hitherto described species are the segments so long as in the
present form. Rhabdometra nigropunctata has the longest of
any ; but here the most posterior are only 3 x 1.
There is, to my mind, no doubt that the species which is
nearest akin to that which I here name Rhabdometra cylindrica
is Fuhrmann’s recently described Rh. nwmida*, and it will be
observed that both come from the same part of the world and
they both infest gallinaceous birds. There are, however,
sufficient differences to warrant specific separation. Thus,
Rh. numida is a small species 60-70 mm., and has not, as
already mentioned, elongated posterior proglottids. Although
the two species agree in possessing a long cirrus-sac, that
of Rh. numida is much the longer, reaching as it does to the
middle of the body. Nothing is said as to the development of
* Res. Swed. Zool. Exp. Egypt, Pt. ii. 27, p. 36, 1909.
AVIAN CESTODES. 879
the uterus; but it is, when fully matured, a lobate sac, and
apparently not strictly cylindrical, as in my species. On the
other hand, the paruterine organ is represented as appearing first,
and the course of the vas deferens is as in my species and not as
in some others. Further, the genital apertures of Rh. numida
are behind the middle line of the proglottid, and the testes come
nearer to surrounding the ovary than in other species, except
Rh. cylindrica. Finally, the generative ducts lie between the
water-vessels and ventral to the nerve-cord. I have observed
the same relationship in 2h. cylindrica.
It is possible, of course, that previous authors have not seen
quite fully-developed examples of the species studied by them.
In view, however, of the published figures of the uterus and
paruterine organ in the several species, | am inclined to doubt
this. For example, Ransom’s figure of that organ in Rh. similis*
shows the “ flowing appearance” of the core of the paruterine,
which I find only in that organ when fully developed (see
text-fig. 8, p. 872). As to other species, it does not appear
that the reticular uterus of Rhabdometra cylindrica, a condition
which precedes its cylindrical final form, has any likeness to
what has been observed in previously described forms. This,
indeed, coupled with the form of the paruterine organ and a
number of minor points, such as the posterior position of the
generative apertures, the position of the coil of the vas deferens,
and the distribution of the testes, seem perhaps after all to
necessitate generic separation. But this I leave for the present.
It must be remembered, however, that the scolex of this species
of Ransom is unknown, and that apart from this character it
is hard to distinguish habdometra from Paruterina.
§ A Note upon Otiditenia eupodotidis Leddard.
After communicating to the Society f my note upon this new
tapeworm from Haupodotis kori, it was suggested to me that it
was possibly identical with or near to a recently described new
genus and species Sphyroncotenia uncinata. I had not at that
time seen Mr. Ransom’s paper =, which only (through his kindness)
reached my hands after my memoir was in the press. I was,
therefore, unable to make any comparisons. A consideration of
the structure of this genus Sphyroncotenia, leads me to revise
what I have written concerning the affinities of Oliditenia ; but
the two genera are not identical. In Sphyroncotenia the body
is much more elongate than in Otiditenia; the genital pores are
unilateral; there are many rows of minute hooks upon the
rostellum, and the uterus appears to be racemose and to extend
* Loc. cit. p. 35, fig. 26.
+ P. Z.S. 1912, p. 194.
+ Ransom, “A New Cestode from an African Bustard,” Proc. U.S. Nat. Mus.
bra a DN 7s Yeu coo
880 DR. F. E. BEDDARD ON
much further into the ripe proglottid than it does in Otiditenia.
I have re-examined my preparations of Otiditenia, and find that
my report upon its structure as regards the above points in which
it differs from Sphyroncotenia is correct as to fact.
But I find that I have missed one point of resemblance to
Sphyroncotenia, and through it, to the subfamily Idiogenine
of the Davaineide, This has, of course, an important bearing
upon the classification and position of Otiditenia in the system.
While admitting its resemblances to Davainea, and by inference
to the Davaineide, I was inclined to place Otiditenia nearer to
Choanotenia and its allies. This was, undoubtedly, due to my
not having seen a paruterine organ, though its presumed
absence was not made use of in the generic definition *, or in the
résumé of the most noteworthy characters of the genus following
the definition. Nor, indeed, do the nearly mature proglottids
show any structure exactly resembling the paruterine organ of
Idiogenes, Stilesia, Anonchotenia, Sphyroncotenia, Rhabdometra,
and other forms as figured by various zoologists im memoirs
known to me. In all of these instances, and in others, the
paruterine organ is represented as a structure of fibrous appear-
ance and of limited size, formed apparently from a metamorphosis
of the medullary ground-tissue in the immediate neighbourhood
of the uterus or from the walls of the uterus itself. This latter
origin is asserted by Gough t for Avitellina centripunctata, while
Ransom’s figure t of a “‘ mature segment becoming gravid” of
Rhabdometra similis may be interpreted ina like manner. But
whatever be the origin of these paruterine organs §—and both
Fuhrmann and Gough believe them to be not strictly homologous
through the series—they would appear to have been described as
small bodies lying in, and possibly derived from, but ultimately
independent of, the medullary parenchyma.
In Otiditenia, however, the more mature segments show an
alteration in the medullary parenchyma to which J have referred,
and which I have figured in my memoir upon that genus ||
This alteration affects the whole of the medullary parenchyma as
seen in that section 4] of a nearly mature proglottid. It is visible
up to the circular muscular layer which forms the line of
demarcation between the cortex and the medulla, except where
it is separated therefrom by the ventral water-vascular tube as is
also shown in my figure. The dorsal smaller water-vascular vessel
lies well within the core of medullary parenchyma, as is also
shown in the figure referred to. There is not, therefore, to be
* Loc. cit. p. 220.
+ “A Monograph of the Tape-Worms of the Subfamily Avitellining.” Quart.
Journ. Micr. Sci. lvi. pt. 2, 1911, p. 375.
+ “The Teenioid Cestodes of N. American Dirds.” Bull. U.S. Nat. Mus. No. 69,
1909, fig. 23, p. 31.
§ I do not refer here to the multiple paruterine organs of Davainea, Zschokkeella,
etc.
\| Loe. cit. p. 218, text-figs. 23, 24, 26, 29.
§| Loe. cit. fig. 29, p. 212)
ae ae
AVIAN CESTODES. 881
observed the formation out of the medullary parenchyma of a
definite structure that can be called a separate organ, since the
modified region of the proglottid extends over the whole medullary
parenchyma and includes the dorsal vascular tube.
J have also figured in my paper referred to stages which are
anterior to that which has just been redescribed. In text-fig. 26
of the paper referred to*, three proglottids somewhat younger
are represented in sagittal section. A glance at this figure
might convey the suggestion that a definite paruterine organ of
limited extent lay in each of these proglottids, narrower at one
end (where the letter “'l’” in the diagram is placed) and wider at
the other. Furthermore, the slightly twisted outline of the
(alleged) paruterine organ recalls that of, for example, Rhabdo-
metra nullicollis t. A more careful scrutiny of these sections,
however, brings to light the following faets which are of importance
in the matter. Although, as depicted in my illustration, the edge
of the (alleged) paruterine organ is apt to be wavy and thus to
create inequalities in its diameter, suggestive of a solid body of
irregularly curved outline, it will be found that the waviness is
closely followed by the layer of transverse musculature which
separates, in this as in other tapeworms, the cortical and medullary
layers. Unequal contraction during preservation is, as I think,
responsible for this undulating disposition of the line of transverse
muscular fibres. The object, however, of my figures referred to
was not to show the structure of the medullary parenchyma but
to indicate the position and relations of the uterus. ‘The minute
structure of the medullary region in this stage is less modified
than that of the older proglottids already referred to. The
medullary groundwork is traversed by numerous rather stout
muscular fibres, running mainly if not entirely in a dorso-ventral
direction. These are very frequent, but are single and not
ageregated into bundles excepting at the anterior end of the
proglottid; here the testes of this segment in front are separated
by a thicker layer of these muscles from the parenchyma of the
ensuing segment, the groundwork is comparatively dense, and
there are abundant nuclei. I have recognised that in those pro-
glottids, as well as in the more mature ones, the dorsal water-
vessels are included in the medullary tissue. In comparing this
stage with the older one that has just been described, it appears
that the latter differs only in the degeneration of the mus-
cular fibres of the ground-tissue, which produces the more
fibrous and, at the same time, laxer appearance of the medullary
parenchyma, which, however, may be more resistant, and which
is still further exaggerated in the distended perfectly ripe and
detachable proglottids at the end of the worm’s body. This laxity
favours the movement into the interior of the embryos from
the uterus, which I have described in my paper as occurring in
* P. 204.
+ Ransom, Bull. U.S. Nat. Mus. No. 69, p. 29, fig. -21, 1909.
882 DR. F. E. BEDDARD ON
these proglottids. It should, furthermore, be noticed that while
the medullary tissue is, as already stated, separated anteriorly
from the testes, there is no such separation posteriorly where it
abuts upon the uterus, nor is the epithelial lining of the latter
apparent in these older ‘proglottids ; thus the transference of the
eggs into the medullary parenchyma is render ed easier. Finally,
I have observed that the calcareous corpuscles tend to accumulate
more thickly where the medullary parenchyma touches the uterus,
though they are also present elsewhere and in some numbers
here and there.
To compare with the above older stages in the growth of the
sheltering apparatus for, the developing embryos, I have again
studied younger stages such as is represented in text-fig. 25 of
my paper referred to*, I have, however, more especially studied
sagittal sections, as in the case of the older proglottids. In such
sections there should be visible the origin of the paruterine body,
were this structure in the genus Oliditenia of the same nature
as that of Rhabdometra, etc. But I can find no trace of any
particular condensation and fibrillation of a definite region of the
medullary parenchyma which might later spread and involve the
whole region, which is thus inv olved in mature proglottids, but
for other reasons.
Not only is there nothing of this kind to be observed in the
less mature proglottids, but there is no approach towards
the fibrous appearance of the mature proglottids. This latter
appearance is, in fact, produced by a new growth, which seems to
be very remarkable. I have already referred in my account of
the proglottids of intermediate age to abundant dorso-ventral
muscular fibres in the medulla; it is, perhaps, to the unequal
contraction of these by the pr eser vative reagent that the irregular
outline of the medullar y parenchyma is due, which I have
commented upon as simulating a paruterine organ of limited
extent, lying in the medulla ary parenchyma. I now eall
attention again, In a more particular way, to these muscle-fibres.
In his account of tapeworms of the subf: amily Avitellinine (of the
family Anoplocephalidze), Gough f enters in some detail into the
histology of Avitellina centripunctata, including that of the
muscular system, which I may conveniently take as a basis of
comparison with Otiditenia, which shows an important difference.
Gough points out that the dorso-ventral muscles which traverse
the medullary parenchyma in that direction, consist mainly of
bipolar myoblasts with terminal fibrille and, to a less degree, of
bipolar myoblasts with lateral fibrille. The larger muscular fibres,
which are tubiform with an axially contained myoblast, are
limited to the longitudinal muscular layer. My own observations
upon various genera which I have examined confirm this
generalisation. The delicate dorso-ventral muscle-fibres of such
* Loe. cit. p. 202.
+ Quart. Journ. Mier. Sci. vol. lvi. 1911, p. 347 et seq.
AVIAN CESTODES. 883
tapeworms as I have known hitherto, often curled into a spiral or
at least an undulating course, ave excessively slender as compared
with the longitudinal muscles. In Otidiiceenia, however, we meet
with the very surprising state of affairs represented in the annexed
figure (text-fig. 11). Imbedded in the nucleated parenchyma
are numerous scattered muscular fibres, which have a general
Text-figure 11.
Sagittal section through a portion of proglottid of Otiditenia ewpodotidis.
1. Longitudinal, ¢. Transverse, d.v. Dorso-ventral muscles.
m. Medullary layer.
dorso-ventral direction and must be regarded as the dorso-ventral
musculature. But the fibres themselves are distinctly of the
character of the fibres of the longitudinal muscular layer. They
are long, straight fibres of even diameter, and I traced them
through the layer of cireular fibres which bounds the medullary
884 DR. F, E. BEDDARD ON
parenchyma externally. These fibres, however, are not exactly
like those which constitute the longitudinal muscular bundles.
Although very wide when compared with the delicate fibrils that
one expects to find in this situation, they are little if anything
more than half the width of the longitudinal fibres. There is
also a difference in the way in which they absorb the logwood
stain, indicating a denser, because more deeply stained, outer
layer. I do not attempt a further description of these fibres, as
the material was not preserved in a way likely to bring out greater
detail of structure. But enough is plainly visible to show that
the dorso-ventral musculature in this worm is different from
that usually met with in this situation among tapeworms. It is
now necessary to point out that, in the younger proglottids, these
thick dorso-ventral muscles are not visible. As the mode of pre-
servation and staining has been identical, there can be no question
here of a failure to detect the fibres in question, which I have
looked for both in transverse and sagittal sections. Fine delicate
fibrils can be seen, but nothing like the muscles just described,
which are even: recognisable under quite low powers. It seems
clear, therefore, that they appear; but whether they do so in the
shape of new fibres or of a thickening of more slender fibres
present in the younger proglottids, | am unable to say. But in
any case there is a chan ge of structure in the medullary paren-
ehyma as it grows older which is not a degeneration, and which
appears, therefore, to be a preparation for some function, which is
possibly that of a receptacle for the ripe embryos. It is for these
reasons that I think myself justified in speaking of a paruterine
organ in Ofiditenia which is, however, of a most generalised
kind.
The above description amounts in reality chiefly to an
emphasised re-assertion of the facts concerning the medullary
parenchyma of Otiditenia, already dealt with in my paper on
that genus. The facts lead me to the inference that we have in
this tapeworm the commencement of the formation of the
paruterine organ, which is more differentiated in other genera of
Davaineide, as well as in the Paruterine among the Hymeno-
lepidide. The paruterine organ of Stilesia, and that of its
immediate ally Avitellina, seems to me to be a different structure
altogether, though serving much the same function. In Oétidi-
tenia, then, there is an alteration of structure in the whole
medullary layer in the direction of increased firmness; but there
is no special part of that parenchyma set apart for the shelteri ing
of the growing embryos. In a sense, therefore, I was right in
not describing the existence of a paruteri ine organ ; but. with
equal truth it may be said that this genus has the equivalent of
a paruterine organ. In this genus we see the next stage to that
exhibited by many genera in which the ripe embryos lie in the
unaltered parenchyma, such as Oochoristica, Linstowia, ete. A.
slight increase of specialisation of the conditions observable in
Otiditenia leads us at once to such a form as Sphyroncotenia,
AVIAN CESTODES. 885
where a large conical paruterine organ exists which is distinct
from the surrounding medullary parenchyma.
§ Systematic position of Otiditenia.
I shall now reconsider the systematic position of Otiditenia in
the light of the foregoing revision of certain facts in its anatomy.
As to the hooks which would form so important a means of
preliminary family identification, I am not yet certain whether
they are or are not the typical Davaineid hooks. They may well
be so; but as I have not been able to view them sideways in my
preparations I am unable to be positive. They may prove to be
like those of Oligorchis paucitesticulatus * for example. There is,
however, no doubt that this genus is not in any case a near ally
of Oligorchis or Hymenolepis, so that we may perhaps fairly
assume that the hooks after all conform to the idea that Otidi-
tenia is to be referred to the Davaineide. Of this family
Mr. Ransom has lately? made a useful table of classification
which is an extension quite up to the present date of the table
in his revision of the Cyclophyllidea £.
From the table in question it appears that Otiditenia will
come nearest to Chapmania. The matter for immediate
settlement is, therefore, whether the two genera are to be
regarded as identical, in which case my name will obviously have
to be dropped. It must be remembered, however, that this near
alliance depends upon whether we are to look upon Otiditenia as
possessing a paruterine organ ; otherwise (still considering it for
other reasons to be a Davaineid) Otiditenia will be nearer to, or
identical with, either Davainea or Ophryocotyle. As to Davainea,
we may at once dismiss the idea of near affinity; for in that
genus the ripe embryos are included in numerous separate
paruterine sacs quite unlike the paruterine organs of Idiogenine.
The knowledge of the genus Ophryocotyle mainly depends upon
the descriptions of O. insignis of Lonnberg, the most recent
of which is in a memoir by Fuhrmann §. This worm is to be
at once distinguished from mine by the immense number of
hooks, 2000, which are disposed in an undulating line round a
particularly large rostellum:; furthermore, the uterus, which is
slightly lobate in form, lies behind the ovary, while the testes
are dorsal to as well as behind the ovary. Moreover, the uterus
shows no tendency to break up; it is conceivable, however, that
it might later, in view of the very late breaking up of the uterus
in Otiditenia. Finally, Ophryocotyle is to be characterised by
multiple rows of minute hooks upon a portion of the suckers.
This latter character does not seem to be found in-Otiditenia.
bE)
* Fuhrmann in “‘Nordische Vogel-Cestcden aus dem Museum von Goteborg,
Medd. Goteborg. Mus. Zool., Afd. i. p. 18, fig. 8.
+ “ A New Cestode from an African Bustard,” Proc. U.S. Nat. Mus. xl. 1911, p. 637.
t Bull. U.S. Nat. Mus. No. 69, 1909.
§ Centralbl. f. Bakt. Paras. Bd. xlix. 1909, p. 94.
886 DR. F. E, BEDDARD ON
But in the case of these hooks upon the suckers, it would appear
that in Chapmania they tend to drop off. This difference,
therefore, between Otiditenia on the one hand, and_ both
Ophr yocot yle and Chapmania on the other hand, must be held
in reserve until more specimens have been examined. There is,
however, I think, no doubt that Otiditenia is quite distinct from
Ophryoc atyle, if only by reason of the characteristic rostellam of
the latter. There now remains only the genus Chapmania. ‘The
first obvious point of difference between the two supposed genera
is the armature of the suckers in Chapmania; but, as alveady
admitted, we cannot apparently dwell too strongly upon this, for
the reason that these hooks are said to be occasionally shed from
the suckers in Chapmania. I haye, however, examined the
suckers in two specimens of Ofiditenia ; and the examination of
two examples lends naturally further support to the view that
the hooks are really missing. Apart from this, there are
apparently two main points of difference which forbid a fusion of
these genera. In Chapmania tapika—which species alone comes
into comparison with Otiditenia, for Ch. taurika has unilateral
generative pores and in other respects differs perhaps to a generic
extent from its supposed congener—a tentacle arises from each
sucker; this is figured by Furhmann as elongated and conical.
I have found nothing of the kind in Jongitudinal and transverse
sections of the scolex of Otiditenia. It may be urged that this
failure to discover a similarity may be due to ‘the complete retrac-
tion of the sucker tentacle, and thus to the difliculty of detecting
it. This may be so; but in the meantime I have seen, in a
tapeworm from Vumida (which may perhaps be the very species,
Chapmania tapika), the tentacle freely moving about in the
living worm. Having thus recognised the structure, it is of
course less likely that I should miss it in examples where it was
ravefully looked for. Besides, the apparent non-retraction of
this tentacle in the preserved examples of Chapmania examined
by Fubrmann, leads to the inference that it would be present in
an unretracted condition in my spirit-preserved specimens of
Otiditenia, were it a character of that species. The next point of
difference is the paruterine organ. If we are to regard the
modified medullary parenchyma in its entirety as the paruterine
organ in Otiditenia, the corresponding organ of Chapmania
as figured by Clerc * is distinctly different.
Neither Fubrmann nor Clere gives much in the way of deserip-
tion of the organ. Judging from the figure the paruterine organ
of Chapmania only occupies about half of the ripe proglottid.
It extends towards the uterus, which occupies about the other
half, and ends on its side turned towards the uterus in a flat
surface. ‘This is obviously totally different from the structure
which I have figured in Otiditenia, and considered to be possibly
a paruterine organ. On the other hand, the breadth of the
* Centralbl. f. Bakt. u. Paras. Bd. xlii. 1906, p. 722.
= eee ee ee ae
AVIAN CESLODES. 887
paruterine organ in Chapmania lends some support to a
comparison ; for it is as wideas the medullary parenchyma, which
it entirely fills anteriorly, thus contrasting. with the much
narrower paruterine organ as figured in Jdiogenes. I should also
add that the supposed paruterine organ of Otiditenia has no
line of demarcation from the uterus such as is figured by Clere in
Chapmania. As points of minor importance, the uterus is lobate
in Chapmania and ends much further forward in the segment
than it does in Ofiditenia. The ripe and detached proglottid
figured by Fuhrmann * is apparently not unlike that of
Otiditenia. But it may be seen that the paruterine organ is
more or less completely filled with the ripe embryos, whereas
in Otiditenia as I have mentioned, the ripe embryos are not
scattered throughout the whole of the supposed paruterine
organ.
The testes of Chapmania are described as being dorsal, whereas
in Otiditenia they are posterior, and no more dorsal than ventral.
Concerning the muscular system of the genus Chapmania, there
is a difference of opinion between Fuhrmann and Clere. The latter
regards it as feebly developed, the former as strong; in the latter
event Otiditenia agrees with Chapmania. One would like to
know something of the genera Ascometra and Schistometra of
Cholodkovsky, which are to me at present merely names, being
included in a Russian catalogue of parasitic worms’. As these
genera occur in Bustards they are quite possibly Davaineids. I
do not attempt to redefine Otiditenia until I learn whether it be
held by others that the paruterine organ described above is a
structure referable to that category, and therefore of great
importance as a generic character among the Davaineide, to
which family I now distinctly refer Oliditenia.
* Res. Swed. Zool. Exp. Egypt, Pt. mi. No. 27, p. 22, fig. 16, 1909.
+ Cf. Zool. Rec. 1912.
ON THE FEET OF THE CANIDZ AND URSIDA. 913
49, On the Feet and other External Features of the Canidee
and? "Ursidese eo ywk. Pocock, FRS2 JPM. Se,
F.Z.S., Curator of Mammals.
[Received May 19, 1914: Read June 9, 1914. |
(Text-figures 1-13.)
INDEX.
Page
The Feet, Rhinaria, and Facial Vibrissee of the Canide ... 913
ClassiicationsongherCanidserenrereeereeeeeereeeee eect ecses een OT
MhewMlecirotaiherWinsidcepemsaseocesen eee ne eee ates) O29
Rhmaningor cheslirsidacwerkee eee teen ee ake | BT
Glassiicatrontottne) Unsidaewee eee eee eee eee eee 88
Family CANID &.
Apart from the feet, the external features dealt with in this
paper are the rhinaria and the facial vibrisse.
The feet of the typical Canide are highly specialised, and show
a close adaptive resemblance to those of the Felidz in the form
of the plantar pad, the strong curvature of the line of the pads
of digits two to five, in the backward position of the first digit of
the fore foot, its absence from the hind foot, the hairiness of the
area behind the plantar * pads, and the persistence of the single
carpal pad on the fore feet. In their structural uniformity within
the families, the feet of the Canidez and Felide show a marked
contrast to those of the Viverride, Mustelide, Procyonide, and
Urside.
In most wild species of Canidz the feet differ only in minor
points from those of domestic breeds already described +. The
four principal digits are united by integument up to the base of
the pads. The webbing thus formed may be wide or narrow, and
clothed with longer or shorter hairs, according to the species. As
in other Fissiped Carnivora, the hair grows in tufts between the
pads of the feet and on the upper side of the webs. It is usually
thickest and longest on the webs, the underside of the digits
themselves showing a naked or nearly naked streak.
The rhinariwm is always large and moist, there being apparently
very little variation with respect to the extent to which the hair
of the muzzle encroaches upon it. The upper lip below it is
always divided by a narrow moist distensible area, which presents
the form of a mere slit when its hairy margins are approximated
in the middle line.
The normal tufts of facial vibrisse are always present, but the
* T use this term indifferently for the main pad of both fore and hind feet.
+ P.Z.S. 1914, pp. 478-484. The method adopted in that case of ascertaining
the extent of the web by cutting the hairs short has been followed in the present
communication dealing with the feet of the wild species of Canidze and with the
Urside. The figures represent the paws with the hairs cut and the digits distended.
914 MR, R. I. POCOCK ON THE FEET
vibrisse vary in length and number according to the species.
The interramal tuft is placed nearly in a vertical line with the
corner of the mouth when closed. As in most Fissiped Carnivores,
there are two genal tufts, one behind the corner of the mouth,
nearly in a vertical line beneath the posterior canthus of the eye,
the other usually much higher up the cheek and farther back.
The genera of Canide hitherto established and admitted rest
mainly upon cranial and dental characters.
Speothos venaticus Lund.
(Text-fig. 1, A, B.)
The feet of this rare dog, of which I have only been able to
see dried skins, were figured by Flower (P. Z.8. 1880, p. 70), who
dismissed them with a mere reference, and this figure was repro-
duced by Mivart in his ‘Monograph of the Canide’ (p. xv.) to
illustrate the structure of the feet characteristic of that family.
As a matter of fact, the feet differ from those of all other genera
of the Canide in two very important particulars, namely, the ex-
tensive basal fusion of the third and fourth digital pads of both
the front and hind feet, and the approximation of the digital
pad of the first digit of the fore foot to the inner proximal
angle of the plantar pad. Moreover, the area between the
large plantar and the carpal pads of the fore foot 1s somewhat
scantily hairy, especially externally, where a nearly naked strip
of integument passes from pad to pad, and the edges of the inter-
digital integument connecting the second and third and the
fourth and fifth digits is naked on both the anterior and the
posterior paws, and the area between the digital and plantar
pads appears to be sparsely covered with short hairs *.
From its low position, the first digit, it seems, must reach the
ground when the animal is standing in the normal position,
especially if the soil be soft. Coupled with the scantiness of the
hairy clothing of the area above or behind the plantar pad, this
suggests that Speothos is more plantigrade than any other existing
dog; and it may be recorded in this connection that the mother
of a specimen sent to the Gardens in 1879 (P. Z. 8. 1879, p. 664)
was killed in a creek and that two of the skins in the British
Museum are labelled ‘‘ Shot while running along creek.”
If this dog habitually haunts the borders of streams, its planti-
grade and scantily hairy feet must be an advantage for progres-
sion on sandy or muddy banks. It appears to me to be impossible
* Except for the forward position of the first digit and the fusion of the third and
fourth digital pads in the fore foot, Flower’s figure does not show these features
well, and it is noticeable that the third and fourth digital pads on the hind foot are
represented as separated throughout. The shape of the pads too and the median
position of the carpal pad throw doubt upon the reliability of the figure. Hence it
may be that the very marked asymmetry between the second and fifth digits on both
feet is also exaggerated. But since the figure was taken from a fresh example that
point may be correct. If so, it is full of interest; but on dried skins I cannot find
convineing evidence that the second digit is so far in advance of the fifth as Flower’s
figure indicates.
Text-figure 1.
OF THE CANID/ AND URSID#. 915
ssSS8xiy
= owt
SS
to decide whether the peculiarity of the feet in the matter of the
low position of the first digit and carpal pad is a secondarily
acquired adaptation to conditions or whether it is a retained
primitive character. I incline to the latter opinion, because these
peculiarities are present in the newly born pups of species of
dogs with normal feet when adult (text-fig. 1, C)*.
* In Lund’s original figure of Sp. venaticus (Kongl. Danske Vid. Selsk. xi.
pl. 61, 1845) the first digit is shown in its correct position, but there is a broad web
between the remaining four, giving a palmate appearance to the spread paw, which
is certainly inaccurate so far as the third and fourth digits are concerned.
B. Right hind foot of same.
C. Right fore foot of newly-born cub of Canis pallipes, showing the low position of the pollex.
A. Right fore foot of Speothos venaticus, from dried skin.
916 MR, RB, I. POCOCK ON THE FEET
Cuon primevus Hodgs.
(Text-fig. 2.)
Apart from Blanford’s brief reference to the feet of this species,
which he described as having “long hair between the foot-pads,”
I am not acquainted with any description of them.
I have only seen the feet of two puppies, about four months
old. In the relative positions of the carpal and plantar pads and
the pad of the first digit, the fore foot resembles that of Canis
anthus tolerably closely, but the carpal pad is longer and more
prominent. Marked differences are noticeable in connection
Text-figure 2.
Wy -
Mite, Lh
ae
¥)
4
Par ',
\\\ ¢ ZEYY)
S1\ \ yy
WY L
NN)
~s
SO
Cuon primevus.
A. Left fore foot.
B. Rhinarium from the front.
C. Side view of face, showing vibrisse and rhinarium.
with the four main digits. The third and fourth are widely
separable, the space between them when distended being equal
to that between the second and third and the fourth and fifth.
The edge of the web joining them is smooth and not sharply
differentiated from them by the hairy covering seen in most —
species of Canide. The edge of the web between the second
and third and fourth and fifth digits, is not naked and forms a
tolerably evenly curved line. The greater part of the sole between
the four digital pads and the broadly cordate plantar pad is
OF THE CANIDE AND URSIDA. 917
scantily hairy, but laterally, close to the proximal margins of the
pads and behind the naked rim above described, the hair grows
in the form of a long fringe. This combination of features
is only found in one other dog that I have examined, namely,
Lycaon pictus.
Similar features are presented by the hind foot, which, how-
ever, 1S thinner and longer than the fore foot, the third and
fourth toes being more prominent and less widely separable
and the plantar pad narrower. z
The rhinariwm is bluntly rounded in profile view. Its inferior
edge seen from the front is strongly angular, the margin below
the nostrils being deep anteriorly and narrow and_ shallow
posteriorly beneath the slit; the nostrils are smallish and widely
spaced, and the median groove does not extend up between
them.
The facial vibrisse are normal in position and of moderate
length, and in the specimen examined the superior genal tuft
consisted of two unusually widely spaced bristles.
Lycaon pictus Temm.
(Text-fig. 3.)
A single example of LZ. pictus sharicus examined.
The main peculiarity about the feet of this dog, namely, the
suppression of the first digit of the fore foot, is well known, but
IT am not aware that other characters have been recorded.
In one feature at least the fore foot recalls that of Vulpes
vulpes, namely, in the length and narrowness of the area between
the plantar and carpal pads, but here the resemblance ceases, for
both these pads are large, as in Canis.
The paw itself is strikingly like that of Cuon, except that the
digits are longer. Very suggestive of kinship between the two
genera are the equality in the spacing of the four toes, due to the
comparatively wide separation between the third and fourth digits,
the nakedness of the edge of the web which joins these two digits
together, and the great length of the hairs fringing the proximal
margin of the digital pads, and the scantiness of the hairs clothing
the sole between the plantar pad and these fringes.
The hind foot is like the front, but is narrower and not so
widely splayed.
The chief interest attaching to the feet of Lycaon is their
likeness in the particulars mentioned to those of Cuon, espe-
cially as kinship between these two has been suggested on other
grounds.
The rhinariwm is large and wide, rounded anteriorly in profile
view ; its upper and lower edges parallel and transverse when
seen from the front, the inferior edge sinvous and without any
marked median inferior prolongation. The nostrils are large,
rounded, and somewhat widely separated, the posterior slit of the
nostril is bordered below at its posterior end by a thick area of
918 MR. R. I, POCOCK ON THE FEET
moist black integument; a somewhat deep median groove ex-
tends between the nostrils upwards from the cleft of the upper
lip. This rhinarium is very different from that of Cuon and the
other species and genera of Canide examined.
Text-figure 3.
~~
. WSG SS
SAX“sxs“cKs
SSWWay
Ss
=
— ~S
Lycaon pictus.
A. Left fore foot. a. Naked margin of web joining third and fourth toes.
B. Lateral view of face, showing rhinarium and vibriss.
C. Rhinarium from the front.
Facial vibrisse short and not numerous, except that the inter-
ramal and inferior genal tufts, which are normal in position,
consist of about four bristles each. The superior genal, consisting
of one bristle only, is set much lower than in other Canide.
Canis anthus Linn.
(Text-fig. 4, A, B.)
A single example of this Jackal from Morocco.
Fore feet short, especially the third and fourth digits, which
OF THE CANIDH AND URSIDZ. 919
are somewhat tightly tied together, the distance between them
much less than that which separates them from the second and
fifth respectively. Digital pads moderately large; plantar pad
large, both long and wide. Carpal pad large, wider than long.
to)
Hind foot with digits less widely spread than those of front
foot, plantar pad much smaller both in width and length.
Text-figure 4.
‘
~
Wn
ETN
ie
Ki
If
Canis anthus and C. mesomelas.
A. Right fore foot of Canis anthus. C. Right fore foot of Canis mesomelas.
B. Right hind foot of same. D. Right hind foot of same.
K. Anterior view of rhinarium of C. mesomelas.
920 MR. R. I. POCOCK ON THE FEET
Canis mesomelas Schreb.
(Text-fig. 4, C—E.)
The fore foot of this species is considerably more “foxy” in
form than that of Canis anthus. 'The area between the carpal
and plantar pads is longer and narrower, the carpal pad is smaller,
the plantar pad is narrower and more overgrown with hairs in
the middle behind; the area between the plantar pad and the
notch between the third and fourth digits is longer, the pads of
these digits are tied more tightly together, and the edge of
the web between them and the lateral digits is more deeply
emarginate.
The hind foot differs from the fore foot in the smallness of the
plantar pad and in the still deeper emargination of the edge of
the lateral web.
The rhinarium is acutely rounded anteriorly in profile; from
the front view its upper edge is straight with obtusely rounded
angles, and its lower edge acutely angled mesially with obliquely
sloping sides, the area beneath the nostrils in front being some-
what shallow and becoming progressively shallower laterally and
posteriorly beneath the narial slit.
The facial vibrisse ave normal in position and moderately
long (see supra, p. 901).
Cerdocyon microtis Sclater.
(Text-fig. 5.)
= Canis sclateri Allen.
A single specimen from the Amazons.
Fore feet longer and more loosely webbed than in C. anthus,
the third and fourth digits joined by a wider web, the distance
between them only a little less than that between the second and
third and fourth and fifth. Plantar pad large, but rather smaller
relatively than in C. anthus, its median lobe wider and rounder.
Carpal pad high up, very small, and conical. Claws short.
Hind feet longer and narrower than fore feet and with smaller
plantar pad, the posterior borders of the pad not deeply emar-
ginate. Hairs between the pads thick, but not specially long.
The rhinariwm is nearly rectangular anteriorly in profile view.
Seen from the front its upper edge is straight and transverse
with rather widely rounded angles ; its inferior edge is strongly and
tolerably evenly convex, owing to the great depth of the portion
below the nostrils in front, the portion below the slit of the
nostrils behind narrow ; hairy area of the lip below the rhinarium
shallow, its median slit continued upwards on to the rhinarium
as a groove which ascends a little higher than the inferior rim of
the nostrils.
The faciai vibrisse moderately long—shorter, that is to say, than
in C. mesomelas, Ps. gracilis, and the species of Vulpes; normal
in position, except that the superior genal tuft, consisting of two
f i.
OF THE CANIDH AND URSIDA. 921
bristles, is set unusually high, nearly on a level with the posterior
canthus of the eye.
Text-figure 5.
Cerdocyon microtis.
A. Right fore foot. C. Rhinarium from the front.
B. Right hind foot. D. Rhinarium from the side.
Points to be noticed in connection with this species, apart from
the remarkably small size of the ears, which are only 27 inches
(56 mm.) long *, are the comparatively wide and nearly even
spacing of the digital pads when stretched, the very small size of
the carpal and pollical pads, the depth of the rhinarium below
the nostrils in front, and the high position of the superior genal
vibrisse.
* Most of the South American dogs have the ears as long relatively as in typical
foxes (Vulpes vulpes), which about equal Cerdocyon microtis im size and have a skull
of about the same length. But, according to Miller, the ears of V. vulpes range from
82 to 98 mm.
The czecum in this example of C. microtis was short and uncoiled as recorded by
Garrod of C. cancrivorus (= C. thous), and there is very little doubt that these
two species are tolerably closely related. On the other hand, in the specimen of
Ps. gracilis from Cordova and in an example of Ps. azaricus from Mar del Plata, the
cecum was longer and coiled, as described by Garrod of C. antarcticus. There are
discrepancies in the accounts of the cecum of C. azare, Mivart stating it to be
straight and Garrod stating it to be coiled. The name azare, however, has been
given to at least two distinct species, one belonging to the thous- or cancrivorus-
group and another to the culpeus-group of South American dogs (see Thomas, Amn.
Mag. Nat. Hist. (8) vol. xiii. p. 345, 1914). No doubt, Mivart and Garrod had
different species under examination, and that Mivart’s determination was probably
correct may be surmised from the fact that true azar belongs to the thous-group.
922
MR. R. 1. POCOCK ON THE FEET
Pseudalopex gracilis Burm. ?*
(Text-fig. 6.)
A half-grown specimen of this or an allied species of the
culpeus-group from Cordova.
Feet long, the small conical carpal pad high above the plantar
pad, the pad of the first digit about on a level with the mid-point
Text-figure 6.
‘Y
Wrt{\y\)
\
Pseudalopex gracilis ?
A. Left fore foot. C. Rhinarium from the front.
B. Left hind foot. D. Rhinarium from the side.
between the two.
Plantar pad long as compared with its width,
narrowly cordate with deeply emarginate posterior border.
digits moderately long, with small pads and unusually distensible,
The
individual is a little uncertain.
* Typical gracilis came from Mendoza. Hence the determination of this young
OF THE CANIDA AND URSIDA. 923
so that the foot is extraordinarily wide when spread to the fullest
extent, the third and fourth capable of being separated until the
inner edges of the pads are in a transverse, almost straight, line
with the border of the web connecting them which is only lightly
emarginate, the width of this web about twice the length of one
of the pads and exceeding the distance between the pads of the
second and third or fourth and fifth digits. Claws long and
slender. The hairs on the lower surface of the foot abundant,
silky,and long, completely concealing the pads when undisturbed.
Hind foot very like the fore foot, but smaller, the posterior
portion of the plantar pad more overgrown with hairs.
The rhinariwm is rectangularly rounded in front in profile ;
from the front view its upper edge is wide and straight with
nearly rectangularly rounded angles, its inferior border is
mesially angular with obliquely sloping, slightly sinuous sides,
the area below the nostrils being moderately deep in front and
narrow posteriorly below the slit; the hairy area of the lower
lip below the rhinarium in front is moderately deep and the
cleft is continued upwards as a shallow groove between the
nostrils.
The facial vibrisse are long and abundant and normal in
position, about six superciliaries, three interramals, and three to
each of the genal tufts. Im an example of Ps. azaricus from
Mar del Plata, the vibrissee resemble those of the species above
identified as Ps. gracilis.
The feet of Ps. gracilis suggest adaptation to desert conditions,
the spread of the digits and the thick hairy clothing preventing
sinking in the sand.
It will be interesting to see in the future to what extent the
feet of the other fox-like South American dogs, suchas Ps. culpeus,
conform to this type, which differs so widely from that of some
of the true foxes of the Old World, like Vulpes and Otocyon.
Alopex lagopus.
(Text-fig. 7.)
A single old female specimen of 4. lagopus spitzbergensis.
The feet differ from those of all the species of Canide
examined in the presence of a distinct thickish ridge of integu-
ment, passing forwards from the median lobe of the plantar pad
to the area behind the point of junction of the third and fourth
digits, and dividing the glandular depression between the digital
and plantar pads into right and left portions. Moreover, the
area of the integument on the proximal side of the digital
pads is thickened and cushion-like, not gradually sloped away, so
that the partially divided depression is bordered in front and
laterally by elevated walls, making it appear deeper than in other
species. The median longitudinal ridge of skin is not so high as
the plantar pad and as the thickening behind the third and fourth
Proc. Zoou, Soc.—1914, No. LXII. 62
924 MR. R. I. POCOCK ON THE FEET
Text-figure 7.
Alopex lagopus.
A. Right fore foot. C. Rhinarvium from the front.
B. Right hind foot. D. Rhinarium from the side.
digits, and, like the thickenings in question, it is clothed with
longish hairs *.
The feet are shorter and more “ dog-like” than in Vulpes and
Otocyon, owing mainly to the third and fourth toes being shorter
and a little more widely separable. The digital pads are small
and the plantar pad is a little wider than in Vulpes and more
overgrown posteriorly, so that it appears more deeply hollowed
out. The carpal pad is nearer the plantar pad, and is small and
irregularly semicircular in outline. Claws very long.
The rhinariwm in profile view is nearly rectangularly pointed,
and much shorter than in any other species examined. From
the front view the summit is mesially nearly flat with widely-
rounded angles; the inferior edge is not strongly angled, the
* Tn the specimen examined, an old female which had been over ten years in the
Gardens, the hair on the integumental thickenings was worn off by walking on
concrete. But the outline of the digital pads could be distinguished by their scaly
pattern, the thickenings being pitted with hair-follicles.
OF THE CANIDA AND URSIDA. 925
nostrils are large and separated by a very narrow septum, and
the groove from the upper lip extends upwards to a point just
above the inferior border of the nostrils.
The vibrisse are normal in position, and the mystacial and
submental bristles are shorter than in Vulpes or Canis mesomelas.
Until Miller published his ‘Catalogue of the Mammals of
Western Europe,’ the Arctic fox was not regarded as generically
distinct from Vulpes, although Kaup and, later, Gray had applied
generic names to it, because of the unsatisfactory nature of the
characters by which it was distinguished. It cannot be claimed
that the diagnostic features employed by Miller are of great
value, as he himself seems prepared to admit. Most of them are
cranial *, the only external feature mentioned being the shorter
and more rounded ear. If such characters, however, be used for
distinguishing ‘“‘ Alopecoid”’ genera, it will be necessary to give
generic status to nearly every species of fox—ausing “‘ species” in its
old sense. Judged, however, by its feet, the Arctic fox is quite
distinguishable generically from Vulpes vulpes and bengalensis
and from Ofocyon, and the same may be said of the rhinarium 7.
In the extent of the area of the sole applied to the ground the
feet of Alopex lagopus show superficial resemblance to those of
Speothos venaticus, and a comparison between the two species
suggests that the structural modification described as fusion of the
pads in the latter is due to the nakedness of the integumental
cushions behind the pads. But since in Speothos the naked areas
regarded as pads are granular throughout, and show no trace of
hair-follicles like the thickened integument behind the pads in
Alopex, the view put forward in this paper, that the pads of
Speothos ave enlarged and fused, appears to be correct.
Vulpes vulpes Linn.
(Text-fig. 8, A.)
Feet long and narrow with comparatively small pads and long
claws. Carpal pad small and set high above the plantar pad ;
and the digital pad of the first digit (poliex) above the middle of
the area between the carpal and plantar pads.
Plantar pad moderately wide, but subcrescentic in form, owing
to the encroachment of the hair over its median portion
posteriorly. Third and fourth digits especially long and tied
together by a narrow web, so that the interval between them,
when extended, is much less than the interval between the second
* One feature mentioned needs restating. It is said that the interorbital region
of the skull is more elevated than in Vulpes, owing to greater inflation of the
frontal sinuses. Asa matter of fact, the frontal sinuses, as Huxley pointed out, are
undeveloped, as in other “ Alopecoid ” skulls, the inflation in question being caused
by the upward extension of the nasal passages.
+ The caudal gland in the specimen examined was larger than in any species of
Canide that I have looked at. It was a hairless patch of very thick glandular skin
about 25mm. long and one-third longer than wide. There was no underfur mixed
with the hairs surrounding it; and, like the feet, it gave out a strong “foxy” smell.
62*
PA MR. R. I. POCOCK ON THE FEET
and third and the third and fourth digits. The length of the
four digits makes the edge of the web connecting the median
and lateral toes appear widely scooped out. The claws are long
and the hair clothing the area between the digital and plantar
pads is thick, but not long*.
Vulpes bengalensis Shaw.
(Text-fig. 8, B—D.)
The feet are similar in a general way to those of V. vulpes, but
the third and fourth digits are, if anything, relatively longer
Text-figure 8.
L y
Mie
ce a
DWN Zz
i Y
) Y
CMMI,
‘ 7
ZN)
ug
? lf) Ye,
SAA 1
SSS AN
\
x
NIN
DENA
AN
Vulpes and Otocyon.
A. Left fore foot of Vulpes vulpes. D. Rhinarium of same, from the side.
B. Left fore foot of Vulpes bengalensis. EK. Rhinarium of Otoeyon megalotis,
C. Rhinarium of same, from the front. from the side.
F. The same, from the front.
* The hairiness of the soles of the feet may vary seasonally in this species, as it
does in some of the northern Canidee. Very likely, too, variation in this respect
will be found between geographical races of V. vulpes.
OF THE CANIDA AND URSIDA. 927
and the second and fifth shorter. The carpal pad is relatively
larger, the posterior region of the plantar pad is less overgrown
with hair, and the hairs arising just behind the digital pads are
exceedingly long and project forwards beneath the pads as far as
the tip of the long slender claws.
The rhinariwm is slightly acutely rounded in profile view
anteriorly ; from the front its upper edge is tolerably evenly
curved and its inferior border only slightly angled mesially, the
area below the nostril being moderately deep in front and narrow
below the slit posteriorly.
The facial vibrisse are long and normal in position.
Otocyon megalotis Desin.
(Text-fig. 8, E, F.)
In all essential respects the feet of this fox, accorded generic
‘ank mainly by reason of its abnormal dentition, agree with
those of Vulpes vulpes and bengalensis, although the third and
fourth digits are a little shorter. The median lobe of the plantar
pad is a little more prominent, especially as compared with that
of V. bengalensis, and its posterior border is emarginate by the
erowth of hairs as in that species. The carpal pad is very small.
The rhinariwm is slightly elevated above and rectangularly
rounded anteriorly in profile view ; seen from the front its upper
edge is straight with obtusely rounded angles and its lower edge
is angular, the portion below the nostrils being shallow and very
narrow laterally and posteriorly below the slit.
The facial vibrisse ave normal in position and moderately long.
CONCLUSION.
Although only a few species of Canidee, compared with the
numbers known, have been described in the foregoing pages,
they fortunately represent the main groups of generic or
subgeneric rank into which the family has been divided. [t is
probable therefore, | think, that the extremes of variation in
the structure of the feet, in the shape of the rhinarium, and the
disposition of the vibrisse have been observed.
It does not appear to me to be likely that the broad distinetion
between Speothos and the rest will be lessened by the examination
of other species—and probably Lycaon and Cuon, and possibly
Alopex, will also remain isolated; but it would be rash to assume
in the present state of our knowledge that the differences in
foot-structure between the species recorded under Pseudalopex,
Oerdocyon, Canis, and Vulpes will hold good in all cases when
the species related to them come to hand for examination.
That, however, remains to be seen.
In the meantime, it may be useful to tabulate the results so
far achieved to show briefly how the species may be grouped,
and to draw attention to the more salient characters presented
by the feet. In the following table, however, | have only made
928 MR. R. I. POCOCK ON THE FEET
use of the fore feet, partly in the interests of brevity, partly because
they supply the best characters, and partly because the features
presented by the vibrissee and rhinarium call for verification on
fresh material, which is unavailable at the present time :—
a. Pad of pollex set low down close to the posterior lateral angle
of the plantar pad; pads of third and fourth digits basally
united; edge of web between median and lateral digits naked ;
area between digital and plantar pads and between carpal and
plantar pads scantily covered with short hair ............ Speothos venaticus.
a’. Pad of pollex, when present, high above plantar pad ; pads of
third and fourth digits separated; edge of webs joing the
median with the lateral digits hairy ; area between digital pads
and plantar pad and between the latter and the carpal pad
mostly thickiy hairy.
b. Edge of web joining third and fourth digits naked, forming a
definite hairless band; a long fringe rising behind this and
extending round the periphery of the foot behind the digital
pads.
ec. Pollex suppressed, feet long, carpal pad high above plantar
pad ane cin Sa < bbe eal. Seek ames Lycaon pictus.
ce’. Pollex retained; feet shorter, carpal pad moderately high
above plantar Pads. .<s.cn2.-c-con-nteerectbecercsesersmssensass CULO POPU GMS:
b’. Edge of web between third and fourth digits hairy.
d. Glandular depression between pads undivided ; integument
proximal to the pads not specially thickened.
e. Third and fourth digits exceedingly widely separable, the
edge of their web when stretched forming a straight line
about twice the length of the pads of either digit; plantar
pad long and narrow .............00600e00:c01eeeee Pseudalopex gracilis.
e’. Third and fourth digits much less widely separable, the
edge of the web between them emarginate when stretched
and less than the length of the pad of either digit.
f. Plantar pad large, its width exceeding the length between
its median lobe and the margin of the web between the
shorter third and fourth digits; claws short.
g. Carpal pad very small, third and fourth digits more
widely separated, the web joining them, when ex-
tended, as wide as that between them and the
Nateralidigiisy s.ieecee selec cstaecs Cerdocyon microtis.
g’. Carpal pad large, third and fourth digits more closely
united; the connecting web narrower than that
between them and the lateral digits... Canis anthus, mesomelas*.
f’. Plantar pad small, its width considerably less than the
distance between its median lobe and the edge of
the narrow web between the long third and fourth
digits; claws long vecereccesscsseeeese Vulpes vulpes and
bengalensis ; Otocyon megalotis.
d’. Glandular depression between digital and plantar pads
divided by a ridge of thick skin stretching forwards from
the median lobe of the plantar pad; integument surround-
ing proximal portion of digital pads much swollen and
cushion-like; feet otherwise almost as under e’ ......... Alopex lagopus.
The above-given analysis of the characters of the feet of the
Canidee emphasises the distinctness. of Speothos from the rest of
the genera, and does not afford support to the affiliation of Speothos
* The feet of domestic breeds of dogs come under this heading. The feet of
some breeds indeed conform very closely to the type seen in Canis anthus (see
P. ZS. 1914, pp. 478-484).
OF THE CANIDA AND URSIDA. 929
with Cuon and Lycaon in the Cuon-group opposed to the Canis-
group containing Canis, Vulpes, and Otocyon*.
On the contrary, the distinctness of Speothos may, | think, be
justifiably expressed by setting the genus aside in a special
subfamily, the Speothoinz, the remaining genera being called
Canine.
There is, as yet, no agreement respecting the number of genera
into which the Canide are divisible, but all recent zoologists are
in accord in admitting Lycaon, Cuon, Canis, Vulpes, Otocyon and
probably Vyctereutes and Urocyon. Both Vulpes and Canis have
been further subdivided into many genera or subgenera.
Thomas, for instance, has recently shown that, apart from
Speothos, the following South American dogs have been
generically named as follows :—Chrysocyon for jubatus; Dasicyon
for antarcticus ; Cerdocyon for thous (= cancrivorus) and brasi-
hiensis (= azare); Pseudalopex for magellanicus, azaricus, etc.,
and Lycalopex for vetulus. At present, however, these genera
are, | believe, merely nominal, in the sense of being undefined.
There will be time enough to discuss their validity when the
distinctive features have been ascertained and stated ; and the
same may be said for such subdivisions of Vulpes as Vennecus and
Zerda. Possibly extended study of the feet and other external
features may help the settlement of this difficult question.
Family Urnsip# .
The subjoined account of the feet and the noses of the
Urside is based upon the examination of examples of the
following species that have died in the Gardens, namely, the Polar
Bear (Thalarctos maritimus), the American Black Bear (Ursus
americanus), the Himalayan Bear (Zremarctos thibetanus), and
the Sloth Bear (Jelursus wrsinus); and secondly, upon observa-
tions on living examples in the Society’s menagerie and upon
dried skins in the British Museum, The four species in question
exhibit the extreme range of variation in the structure of the
feet, the Polar Bear and the Sloth Bear being at opposite poles
in the matter of modification; and all the other existing species
of bears agreeing, apart from minor details, either with
U. americanus or with 7’. thibetanus.
The general shape of the feet of bears is well known.
Measured from the carpus or the tarsus to the tips of the
phalanges, they are remarkably short and broad. The five digital
pads form a slightly curved line, the second, third, and fourth
being nearly on a level and a little in advance of the first and
fifth. A short distance behind these pads, and separated from
them by a comparatively thin-skinned depression, comes the wide
* By Matthew and Osborn (‘The Age of Mammals,’ 1910). I do not, however,
know the nature of the evidence on which this classification was based.
+ Ann. Mag. Nat. Hist. (8) vol. xiii. p. 352 (1914).
+ For the generic terms adopted for this family see infra, pp. 9389-940.
930 MR. R. I. POCOCK ON THE FEET
and flattish main or plantar pad*. Behind the plantar pad of
the fore foot there is always one additional carpal pad on the
external or ulnar side of the carpus; and on the hind foot there
is always a larger or smaller naked area, which may involve the
whole of the posterior portion of the sole as far back as the heel.
It is mainly, however, in the degree of hairiness of this area
behind the plantar pads of both fore and hind feet that the
greatest variation is exhibited.
Feet of the Polar Bear (Thalarctos maritimus).
(Text-fig. 9, A, B.)
Tn two newly-born cubs of Polar Bears from Spitzbergen, the
digital pads are not webbed, but are separated to the base as in
all bears except Meluwrsus. The depressed area between them
and the very short and wide plantar pad is scantily covered with
very short hairs, and the area behind the plantar pad is similarly
covered, except for the small external carpal pad on the fore foot
and a corresponding, elongated, somewhat piriform, anteriorly
pointed, small, flat pad on the sole of the hind foot, which are
quite naked.
In our adult living examples the soles of the feet, apart from
the digital and plantar pads, the carpal pad, and the corresponding
elliptical area on the hind foot, are thickly covered mostly with
long hair, except the sole of the hind foot, where the hair is
worn short; and in a male specimen a narrow strip of naked
skin extends forwards from the naked elliptical area to the
plantar pad of the hind foot. The feet, in fact, agree with the
description of the feet of the Polar Bear, recently published by
G.S. Miller t, who says :—-“ Fore feet with palmar tubercles and
balls of toes essentially as in U. aretos, but smaller; pad on hind
foot without backward continuation along inner [outer] portion
of sole.” Since Miller did not detect the little naked pad behind
the plantar pad on the hind foot, it is possible that this pad is
sometimes, perhaps seasonally, covered with hair. But its
presence in this newly-born cub is full of significance.
Feet of the black Bear (Ursus americanus).
(Text-figs. 10 and 13, D, F.)
The fore feet of an adult male Black Bear from Newfound-
land agree in essential points with those of the Polar Bear, that
is to say, the digital pads are separated, the depression behind
them is covered thickly with long hairs, and the area behind the
plantar pad is similarly clothed with hairs, from which the carpal
* It is the custom sometimes to call the main pad of the fore foot the “ palmar”
and that of the hind foot the “plantar” pad. But in this paper I have used the
term plantar for the main pads of both fore and hind feet.
+ Cat. Mamm. Western Europe, p. 298,1912. In his description of the hind foot
of this species, as of U. arctos, Miller wrote “inner” for “ outer.”
OF THE CANIDA! AND URSIDE. 93]
Text-figure 9.
Sat ee
Swe SS
Zz
Le,
a
BZ
Z
Newly-born cubs of Polar Bear (Thalarctos maritimus) and of European
Brown Bear (Ursus arctos).
A. Left hind foot of Polar Bear (Thalarctos C. Right hind foot of Brown Bear
maritimus), the digits spread. (Ursus arctos).
B. Left fore foot of same. D. Left fore foot of same.
1 and 5, first and fifth digits ; Cp., carpal pad; p., pad on sole of
hind foot of 7. maritimus.
pad rises like an island. But this pad, the digital pads, and
especially the plantar pad are relatively large. ‘The hind foot,
however, is very different, in being mostly naked almost back to
the heel. On the inner or hallucal side, however, the hair grows
inwards from the edge of the foot for a short distance behind
932 MR. R. I. POCOCK ON THE FEET
the plantar pad, filling up the depression which at this point
separates this pad from the posterior naked part of the sole.
Text-figure 10.
Kira el) HI
/
B
Ursus americanus.
A. Right hind foot. B. Right fore foot.
1 and 5, first and fifth digits; Cp., carpal pad.
Feet of the Himalayan Bear (Tremarctos thibetanus).
(Text-figs. 11 and 13, C.)
The fore feet differ markedly in one or twg points from those
of the Black Bear. 'The area between the digital pads and the
plantar pad is hairy only behind the pads of the second, third,
and fourth digits; behind the first and fifth of these pads it is
naked and the digital pad of the first is smaller and set still _
farther back, its distal end scarcely reaching the proximal end
of that of the second. The plantar pad is large. Behind it
there is a naked depression of thinner skin and the carpal region
is also wholly naked, the ulnar carpal pad forming a large pro-
tuberance and the radial a smaller one. ‘This smooth carpal area
OF THE CANIDA AND URSIDA. 933
is sharply circumseribed behind by the dense clothing of hair
covering the lower side of the leg.
The hind foot broadly resembles that of UJ. americanus, except
that the depression behind the digital pads is clothed with hairs
in the same way as the fore feet and the hairs from the inner
edge do not encroach upon the sole in the depression marking off
the plantar pad postero-internally. The tips of the pads of the
first and fifth digits slightly overlap the proximal ends of those
of the second and fourth respectively.
Text-figure 11.
Wij,
SUC Ba UN mG Ni
Ny Hit
Win
y
sy)
Zaman
SENSORS
aX
eee,
Tremarctos thibetanus.
A. Right hind foot. Bb. Right fore foot.
1 and 5, first and fifth digits ; Cp., carpal pad.
Feet of the Sloth Bear (Melursus ursinus).
(Text-figs. 12 & 13, A, E.)
The feet differ from those of all other species of Ursidee, in
that the digital pads are fused almost to their distal ends,
so that no hair projects between them from the sides of the
digits, and the first and fifth digits are set far forwards as
compared, at all events, with 7’, thibetanus. Moreover, the
depression behind the the digital pads is quite naked, as Gray
pointed out. In the fore feet the carpal area is naked, as in
934 MR. R. I. POCOCK ON THE FEET
T. thibetanus, but this naked area is not sharply defined behind
by a coating of thick hair, but passes insensibly into the skin of
the posterior surface of the leg, which inferiorly is scantily
clothed with short hairs. The sole of the hind foot is entirely
naked, the posterior limit of the plantar pad being marked by
a transverse groove, which expands into a shallow depression on
the hallucal or inner side.
Text-figure 12.
7
2 itty o) 4
awh,
a i! With am
WAG
Melursus ursinus.
C. Left hind foot (too narrow for its length). D. Left fore foot.
1 and 5, first and fifth digits; Cp., carpal pad.
Feet of other Species of Bears.
The feet of the Polar Bear and of the Sloth Bear are unique
in the family, but those of other species, of which I have only
seen dried skins or living specimens, agree in the main with those
either of 7. thibetanus or of U. americanus, U. arctos and
horribilis falling into the same category as U. americanus and
Helarctos malayanus, and 7’. ornatus into that of 7’. thibetanus.
OF THE CANIDA AND URSIDA. 935
G. 8. Miller * describes the feet of the Brown Bear of Western
Europe as follows :—“‘ Balls of the digits [of fore feet] large,
pad-like ... first digit with anterior edge of ball extending about
to middle of that of second, the interval greater than in the case
of the other digit .... main pad wider than long, covering more
than half the surface of the palm, its outer border about twice as
long as its inner, its posterior border slightly concave, its inner
portion at base of thumb [1st digit] marked off from the rest by
a slight furrow; region between main pad and balls of digits
densely furred; wrist-pad about as large as ball of digits, near
outer ulnar margin of palm, its long diameter transverse ; region
between wrist-pad and main pad densely furred... Hind foot
longer than fore foot, pad like that of fore foot, but with a broad
backward extension passing along inner [outer] side nearly or
quite to heel; region between pad and balls of toes and at outer
[inner | side of backward extension densely furred.”
In a newly-born cub of U. arctos (text-fig. 9, C, D, p. 931), the
area between the digital and plantar pads of both fore and hind
feet is scantily covered with very short hairs, the posterior or
heel pad of the hind foot is naked, and there is a well-marked
depression of wrinkled skin on the inner side of the foot between
this pad and the plantar pad. In the fore foot the area behind
the plantar pad is scantily clothed with short hairs, and the conical
carpal pad is situated near the postero-external portion of this
area.
_ So faras I have been able to examine them, the living bears
of this species in the Gardens, namely examples from the White
Sea, Caucasus, Himalayas, Behring Sea, and Alaska, have the
feet as above described by Miller, except that the entire sole of
the hind foot is generally naked, there being usually no extension
of the hair behind the plantar pad on the inner side. In some
Brown Bears, too, there is a narrow strip of scantily-haired skin
extending from the carpal pad to the plantar pad of the fore foot,
and sometimes a small naked area marks the position of a radial
carpal pad. These points may be worth further investigation
from the systematic point of view.
In one of two Grizzly Bears (U. horribilis) from Montana, the
feet seem to resemble those of our Brown Bears; and Mx. Seton’s
figure of the paws of the Grizzly show the same conformity to
the Brown Bear type.
The chief difference between the feet of U. americanus, on the
one hand, and WY. arctos and horribilis, on the other, is that in
the former the first digit and the carpal pad seem to be set
farther back.
Tremarctos thibetanus ranges from Baluchistan to Kastern Asia,
and is represented in Japan by 7’. japonicus, which is probably
* Cat. Mamm. Western Europe, p. 287, 1912.
+ The sole is continuous along the outer, not along the inner side of the foot.
The hairy ingrowth interrupting the continuity of the sole oceurs on the inner or
hallucal side.
936 MR. R. I. POCOCK ON THE FEET
only a subspecies of it. At all events, the feet are the same in
the two forms.
Moreover, the feet of the Malayan Bear (Helarctos malayanus)
agree in all essential respects with those of 7. thibetanus, except
that the hairs in the depression behind the second, third, and
fourth digital pads are much fewer in number, the integument
being scantily furred, and thus approximating the naked con-
dition of this area seen in Melursus. The Andean Bear (7.
ornatus) also resembles 7’. thibetanus in the structure of its feet,
except that the depression behind the digital pads is continuously
and thickly hairy, even behind the pads of the first and fifth
digits.
Structural Adaptation of the Feet to Habits.
The bionomical reason for the differences in the structure of
the feet of existing Urside is obscure. The first thing to note
is the rough correspondence between the hairiness and nakedness
of the sole and the geographical latitudes inhabited by the
species. The hairiest feet of all are found in the Arctic species
(Thalarctos maritimus), and this feature is always assumed,
probably correctly, to be a modification to obviate the likelihood
of slipping on ice. But it must also be remembered that the
haunts of this bear are treeless, and that this species is unable to
climb. South of the range of the Polar Bear come the various
races of Ursus arctos, of U. horribilis, and of U. americanus,
which have larger pads on the fore and hind feet than in
Thalarctos maritimus and the greater part, at all events, of the
sole of the hind feet naked. Even the northern form of these
species, by reason of their hibernation, are never abroad for any
length of time when the cold is severe enough to cover the
ground for weeks at a time under a continuous sheet of frozen
snow. The countries they inhabit are forested, and both the
Black and the Brown Bears are known to climb trees, The
Grizzly does not climb—at all events, asa rule,—but he probably
could do so, if necessary, though, on account of his greater bulk,
not with such ease as the Black Bear and smaller representatives
of the Brown Bear. Nevertheless, neither the Black Bear nor
the Brown Bear seems to be so apt at climbing as the three
naked-footed bears of the Old World, namely, the Himalayan,
the Malayan, and the Sloth *; and living examples of these
species have an obviously clumsier, more shufHing gait than the
northern species, and this awkwardness of movement is due very
largely, if not wholly, to the natural mturning of the fore feet.
* I know nothing of the Andean Bear (U. ornatus) in this connection. Of the
Himalayan Brown Bear (U. arctos isabellinus), Blanford says :—“ They can climb
trees, but, in the Himalayas, at all events, rarely do so” ; of the Himalayan Bear, “it
is more in the habit of climbing trees for fruit [than the Brown Bear], and is not
infrequently found in fruit trees,” and of the Malayan species “this bear is a purely
forest animal and an admirable climber,’ while his account of the Sloth Bear
contains many references to its scansorial habits.
OF THE CANID® AND URSIDA, 937
This characteristic of the limbs is more marked in the Malayan
and the Sloth Bear than in the Himalayan, which in this, as in
other respects, comes nearer the group of bears typified by
U. arctos. I think it probable that the nakedness of the carpal
area of the underside of the fore foot in those three species as
well as the inturning of the paws are adaptations to climbing,
because naked roughish integument will give a better hold on
bark than integument nonenedl with hairs, a Aad during the ascent
or backward descent of a vertical tree-trunk—bears : always climb
down rear end foremost—the upward turn of the fore paws gives
the claws a securer grip on the bark, because their points are set
at right angles to the axis of the trunk, without interfering with
the clasping action of the limb.
Noses of Bears.
The rhinarium of Bears is always large and naked, and is
circumscribed above and at the sides by the short hairs of the
muzzle and upper lips. Usually the hair on the summit of
the muzzle forms nearly a straight line, passing from the posterior
notch of one nostril to that of the other; and beneath the
rhinarium the hairs of the upper lip extend almost or quite to —
the middle line, leaving at most a narrow strip of naked integu-
ment below the rhinarium. I have not been able to examine
sufficiently closely a large enough number of specimens to show
the variation in width to which this strip of integument is liable
in Ursus arctos, horribilis, americanus, Tremarctos thibetanus,
ornatus, but in all these species, as in- Thalarctos maritimus, it
is at most a few millimetres wide, narrower, that is to say, than
the median area of the remem between the § inner edges of the
nostrils.
But in the Sloth Bear (Welursus) the rhinarium 1s very large.
Dorsally it extends forwards so as to overhang the nostrils and
backwards some distance behind the posterior end of the slit of
the nostrils. It is also much wider beneath the slit laterally and
there is a very wide median area of moist skin annexed to the
rhinavium on the upper lip. ‘The only bear possessing a rhinarivin
approaching that of d/elursus in relative size is Helarctos malay-
Gaius, which, in this respect, 1s intermediate between 7'remarctos
thibetanuws iad Melursus ursinus. In both these species the
vreater size of the rhinarium and of the moist naked area below
it, is associated with the mobility of the snout and upper lip,
which is a marked feature in /Helarctos malayanus and reaches
an extreme in Jelursus wrsinus.
The facial vibrisse of Bears are reduced in number and length,
as compared with those of most other Carnivores. A few buccal
and supevciliary bristles are retained, but the genals and inter-
ramals appear to be suppressed as a rule. The genals [ did not
find in any of the dead specimens examined, but in the example
938 MR. R. I. POCOCK ON THE FEET
of 7. thibetanus the interramal tuft was represented by a single
longish hair.
Text-figure 13.
NS\\\ \T
\ SS \M}}
NYG Wy
Lug A sers
\
enti:
Nae a
~
Rhinaria of Bears.
A. Anterior view of rhinarium of Melursus wrsinus.
Bb. Ditto Helarctos malayanus (from a dried skin).
(OF Ditto Tremarctos thibetanus.
D. Ditto Ursus americanus.
Ii. Side view of lips and rhinarium of Welursus wrsinus.
F. Ditto Ursus americanus.
Systematic Value of the Feet.
From time to time the Urside have been split up into a
considerable number of genera and subgenera, based partly upon
external, but mainly upon dental and cranial characters, the only
well-marked species which has never apparently received a special
OF THE CANIDAH AND URSID_E. 939
title, even from Gray, beike U. thibetanus. For instance, we
have Thalarctos (usually altered to Thalassarctos) for mar itimus ;
Ursus for arctos and its allies, Danis for horribilis, Huarctos for
americanus, Tremarctos for ornatus, Helarctos for malayannus: and
Melursus for ursinus. Melursus seems to be admitted on all
hands as valid; but probably no two existing zoologists could be
found to agree stort the others, though a majority would most
likely favour the severance of patireios from Ursus. Flower
and Lydekker (* Mammalia Living and Extinct,’ pp. 558-560,
1891), for example, gave full generic value to Melina g
Ursus, and divided the lait. into the Thalarctine section for
maritimus ; the Ursine for arctos, horribilis, americanus, thibetanus,
ornatus and their allies, and the elaxetine section for malayonus.
Max Weber (Die Stiug. p- 539, 1904) admitted Ursus, with
Thalarctos as a subgenus, JHellerwetion and Melursus, hat only
diagnosed the latter; and Beddard (* Mammalia,’ pp. 442-443)
allowed Ursus and Melursus, dismissing Vhalarctos as a “quite
unnecessary ” genus.
Trouessart (Cat. Mamm. Suppl. pp. 178-182, 1904) followed
Flower and Lydekker in the main, but gave subgeneric value to
the sections of Ur ‘sus, adding Euarctos ie them, aad accorded full
generic status to Tnemacinting | for the 8. Aumentieate Bears.
Finally, Matthew and Osborn (‘The Age of Mammals,’ p- 530,
1910) adopted the four genera, Ursus for the Grizzly, Brown, and
American Black Bears, : and, I presume, for the Himalayan and
Malayan as well, Talanaios for the Polar Bear, 7vemarctos for
the Andean or Spectacled Bear, and Melursus for the Sloth
Bear.
The divergence of opinion with respect to the status of such
species as maritumus, americanus, ornatus, and malayanus, in
dicated by these classifications, suggests that the characters used
for their elevation to the elk oo genera or subgenera cannot be
very well marked *. But in view a the conclusions arising from
the facts established in this paper, the one inter esting point ABOUE
which these authors seem to be in accord, differ as they may
about the four species just quoted, is that thihetanus is inseparable
from Ursus, even in the most restricted sense assigned to that
term.
Beyond stating that the soles of the feet are more hairy in
the Polar Bear, ‘the authors above quoted made no systematic
use of the extremities, although Gray had previously pointed out
some distinguishing featur es presented by them. He detected,
* The classifications of 'Frouessart and of Osborn & Matthews are, however. mere
lists of names, no reasons for the arrangement adopted being given. It would he
interesting to know why these authors, alone of those quoted, give full generic value
to ormatus.
+ Considering the wide field covered by his work, - Gray was head and
shoulders in front of many of his predecessors and successors as a sy stematist in the
strict sense of the word. One is tuo apt to allow his mistakes, arising trom his
curious limitations, to obscure one’s regard for the perspicacity he undoubtedly
possessed in the detection of structural differences,
Proc. Zoon. Soc.—1914, No. LXIII. 63
940 MR. R. I. POCOCK ON THE FEET
for instance, that the area behind the digital pads in the Sloth
Bear is naked, whereas it is hairy in others. But he does not
appear to have noticed the marked differences presented by the
carpal area in the matter of hairiness and nakedness in various
species, and his statement that in 7 halassarctos the *‘ soles of the
feet are hairy with a few callous pads, whereas in Ursus, Helarctos,
and Melursus they are bald and callous,” gives very little idea of
the true state of affairs (see Cat. Carn. ete. Mammalia, pp. 217-
237, 1869).
Judged by the characters discussed in this paper, the following
genera seem to me worthy of admission—Velursus, Helarctos,
Tremarctos, Ursus, and Thalarctos. They may be defined as
follows :—
a. Digital pads fused almost up to their distal ends, depression
between them and the plantar pad of both fore and hind feet
naked; carpal area of fore paw naked with large rounded
external and smaller internal pads; integument of fore leg
behind carpal area scantily covered with short hairs; snout
highly mobile, rhinarium very large, extending to edge of lip
as a broad moist area and overhanging the nostrils above ...... Melursus.
6. Digital pads separated throughout their length, depression
between them and the plantar pads more or less hairy;
integument behind carpal area thickly covered with hair;
snout less mobile, rhinarium smaller and not overhanging the
nostrils.
ec. Carpal area as in Melwrsus, naked and furnished with a
larger rounded external and a smaller internal pad.
d, Hair on upper lip not extending beneath the nostrils in
front, but leaving a comparatively wide moist median
area continuous with the rhinarium above ................... Helarctos*.
d’, Hair on upper lip extending nearly to middle line and
leaving only a narrow naked strip of skin continuous
Wahd ATI NO eeVUREOLUINy. RUA SSH ARN GReUREaR ahcearsbsnesnciaeraesuiice, | LhiAlasHeuhaner:
ce’. Carpal area behind plantar pad thickly hairy, carpal pads
represented by a single rounded eminence on the outer side,
as in the Canidew and Felidae, and sometimes by a smaller
one as well on the inner side ; rhinarium approximately as
in Tremarctos.
«. Pads large as in the preceding genera; sole of hind foot
behind the plantar pad naked, except sometimes for an
ingrowth of hair internally behind the plantar pad......... Ursus.
e’. Pads smaller; sole’ of hind foot behind plantar pad over-
erown with hair except for a small naked flat pad near
the external border ceanisscbespermcee PLLC Chose
Melursus, Helarctos, and Thalarctos ave monotypical. T'vrem-
arctos contains two well-defined species, namely thibetanus and
ornatus (type), which I cannot distinguish externally by any
characters of generic value in my opinion. Nevertheless, the
difference in the smoothness of the integument behind the first
and fifth digital pads in thibetanus and its hairiness in ornatus is
very curious. Ursus contains a doubtful number of species and
subspecies, but I am not acquainted with any external features
* This genus, or subgenus, is usually defined by the shortness and breadth of the
skull, smallness of the ears, length of the tongue, etc. ]
——
OF THE CANIDE AND URSID. 94]
justifying the admission of Danis (type horribilis) and Luarctos
(type americanus), unless the more backward position of the first
digit in the fore paw of americanus and the higher position of
the carpal pad be given generic value—in my opinion, an
exaggerated view of their importance*.
As regards the genealogical position of the genera judged from
their feet, analogy justifies the opinion that the naked-footed
forms with free digital pads, like Helarctos and Tremarctos, ave
the more primitive 7. From a stock probably resembling these
in the particulars named, Melwrsus is specialised on one side by
the fusion of the digital pads and Ursus on another side by the
growth of hair over the carpal region. Thalarctos appears to me
to be nothing but a specialised type of Ursus, adapted for
swimming and movement on ice, its longer and more powerful
canine teeth being developed for the seizing and slaying of
seals.
* Merriam admits Evarctos as a subgenus of Ursus on account of certain cranial
and dental differences. Danis, however, appears to be undefinable (Proc. Biol. Soc.
Wash. x. pp. 65, 83, 1896).
+ In the bears themselves this view finds support in the scantiness and shortness
of the hairs clothing the areas behind the digital and carpal pads in the newly-born
cubs of Thalarctos maritimus and Ursus arctos.
ON MITOSIS IN ENAMEL CELLS. 943
EXHIBITIONS AND NOTICES.
June 9, 1914.
Prof. E. A. Mincutn, M.A., F.R.S., Vice-President,
in the Chair.
The Secrerary read the following report on the additions to
the Society’s Menagerie during the month of May 1914 :—
The number of registered additions to the Society’s Menagerie
during the month of May was 230. Of these 116 were
acquired by presentation, 36 by purchase, 43 were received on
deposit, 10 in exchange, and 25 were born in the Gardens.
The number of departures during the same period, by death
and removals, was 220.
Se)
Amongst the additions special attention may be directed to :-—
2 Bladder-nosed Seals (Cystophora cristata) 3 2, from Green-
land, purchased on May 18th.
1 Reindeer (Langifer tarandus) g, born in the Menagerie on
May 9th.
6 Long-tailed Bats (Rhinopoma microphyllum), from India,
new to the Collection, received in exchange on May 12th.
4 Red-headed Bulltinches (Pyrrhala erythrocephala), from the
Himalayas, new to the Collection, and 2 Cotton-Teal (Wettopus
coromandelianus), from India, presented by Alfred Ezra, F.Z.S.,
on May 15th.
2 Cuvier’s Toucans (Ramphastos cuvieri), from the Upper
Amazons, purchased on May 12th.
1 Maximilian’s Parrot (Pionws maaimiliant), from Brazil, and
2 Petz’s Conures (Conurus canicularis), from Mexico, presented
by the Marquess of Tavistock, F.Z.8., on May 28th.
2 Open-bills (Anastomus oscitans), from India, purchased on
May 20th.
Mr. J. Tuornron Carrer, F.Z.S., exhibited microphotographs
showing phases of mitosis in the cells of the enamel organ in
Dasyurus viverrinus and Trichosurus vulpecula. The demonstra-
tion of mitosis in the enamel cells has not been recorded
previously—in fact, little has been published dealing with the
cytological changes which produce the differentiation of the
various cells composing an enamel organ.
In a paper to be submitted to the Society during the next
session, Mr. Carter deals at length with the cytology of the cells
of the enamel organ in Mammals, Reptiles, Fishes, ete., in all of
which abundant evidence of mitosis has been found.
QAd ON A GAZELLE FROM SENNAR.
Mr. R. I. Pocock, F.R.S., F.L.S., F.Z.S., Curator of Mammals,
exhibited on behalf of Major C. P. BrapsHaw an interesting
example of Sémmering’s Gazelle (G@. semmeringi), shot on the
Dinder River in Sennar, and mounted by Messrs. Edward
Gerrard & Sons. The antelope was remarkable for the whiteness
of its pelage, which showed scarcely a trace of the gazelline tint
characteristic of the typical form. That the specimen was not
an albino was shown by the persistence of the typical black
markings on the face, the black horns and hoofs, and the black
tuft on the tail.
NOTICE.
Euvallentinia nom. n. for Vallentinia Stebbing.
Mr. Edward 'T. Browne having kindly called my attention to
the cireumstance that a medusoid genus was named Vallentinia
by him in 1902, and further that a genus of Copepoda was so
named by Norman and Scott in 1906 (Crustacea of Devon and
Cornwall, p. 172), I now propose the new name Huvallentinia for
the genus of Isopoda which I myself have recently called Vallen-
tinia (P. Z.8. 1914, p. 351), leaving it to the joint authors to
propose an alternative name for their genus of Copepoda, unless
it should prove that the want has been already supplied, either
intentionally or incidentally, by some other authority.
(Signed) T. R. R. STesBrNa,
Ephraim Lodge,
The Common,
Tunbridge Wells.
August, 1914.
;
:
No. 185.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.*
June 9th, 1914.
Prof. K. A. Miycuin, M.A., F.R.S., Vice-President,
in the Chair.
The Minutes of the last Scientific Meeting were confirmed.
The Secretary read a Report on the Additions to the Society’s
Menagerie during the month of May, 1914,
Mr. J. THonnron Carrer, F.Z.S., exhibited microphotographs
showing phases of mitosis in the cells of the Enamel Organ in
Dasyurus viverrinus and Trichosurus vulpecula. The demonstra-
tion of mitosis in the enamel cells has not been recorded
previously—in fact, little has been published dealing with the
cytological changes which produce the differentiation of the
various cells composing an enamel organ.
In a paper to be submitted to the Society during the next
session, Mr, Carter deals at length with the cytology of the cells
of the enamel organ in Mammals, Reptiles, Fishes, ete., in all of
which abundant evidence of mitosis has been found.
Mr. R. I, Pococx, F.R.S., F.L.S8., F.Z.8., Curator of Mammals,
exhibited on behalf of Major C. P. BrapsHaw an interesting
example of Sdmmering’s Gazelle (G. semmeringi), shot on the
Dinder River in Sennar, and mounted by Messrs, Edward
Gerrard & Sons, The antelope was remarkable for the whiteness
of its pelage, which showed scarcely a trace of the gazelline tint
characteristic of the typical form. That the specimen was not
an albino was shown by the persistence of the typical black
markings on the face, the black horns and hoofs, and the black
tuft on the tail,
* This Abstract is published by the Sceiety at its offices, Zoological Gardeus,
Regent’s Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications; but it may be obtained on the
day of publication at the price of Sixpence, or, if desired, sent post-free for
the sum of Sia Shillings per annum, payable in adyance.
46
Prof. J. Srantey Garprner, M.A., F.R.S., F.Z.S., gave an
account of a Report on the Fauna of the Monte Bello Islands by
Mr. P. D. Montague, B.A.
The islands are briefly described before considering the fauna ;
they are barren limestone with a limited vegetation and some
mangroves. There are two indigenous mammals, Lagorchestes
conspicillatus (Gould) and Isoodon barrowensis Thom. Of the
twelve land-birds three new forms have been described elsewhere ;
the reptiles number eleven, one being new, and show a marked
reduction in size as compared with continental specimens. Of
insects there are recorded 22 Lepidoptera, 11 Coleoptera, 13
Hymenoptera, and some Orthoptera, ete. Myriapoda number 9
species; there is 1 scorpion. 9 fishes from lagoons are named.
The collections prove conclusively the entire dependence of
these islands for their fanna on the neighbouring continent.
Partial depopulations of the islands owing to drought are sug-
gested, suceeeded by repopulations by means of wind-borne forms
from the south.
Three other papers, also dealing with collections made by
Mr. Montague at the Monte Bello Islands, were received from
Mr. G. GC. Rossox on Cephalopoda, from Miss M. J. Rarasun
on Stalk-eyed Crustaceans, and from Mr. Tom IrepaLe on
Mollusca.
Dr. W. A. Cunnineton, M.A., F.Z.S., read a paper on the
parasitic Eucopepoda collected by the Third Tanganyika Expe-
dition in 1904-1905. The collection consisted of a very small
number of specimens, these forms being evidently much rarer
than the Argulide, which are also external parasitic Copepods
infesting fish. The specimens were referred to the well-known
fresh-water genus Lerneocera, and belong to two species both
described as new. One of these, however, differs considerably
from the more typical members of the genus, and may merit a
separation from it in the light of further knowledge. A third
species of Lerneocera, which was taken on a Nile fish and belongs
to the collections of the British Museum, was also described in
the paper.
Dr. F. E. Bepparp, M.A., F.R.S., F.Z.S., Prosector to the
Society, read a paper containing the description of a new species
of Avian Cestodes and a further discussion of the paruterine organ
in Otiditenia.
Mr. R. I. Pococx, F.R.S., F.LS., F.Z.8., Curator of Mammals,
read a paper “On the Facial Vibrisse of Mammalia,” and pointed
out that in all the principal orders of the class, with one or two
exceptions, the following groups of vibrisse are present in some
genera :—mystaciale on the upper lip, submental on the chin and
lower lip, superciliary over the eyes, gonal on the cheeks, and
interramal on the throat behind the symphysis of the jaw.
47
Within the limits of the orders these tufts are present in the
primitive genera, but more or fewer of them may be lost in the
more specialised types. This fact, coupled with their prevalence
in widely different types, points to the arrangement of the
vibrisse above indicated being exceedingly primitive. The
different modifications of the vibrisse met with in various orders
were briefly pointed out.
A second paper by Mr. Pocock “ On the Feet and other
External Features of the Canide and Urside,” based like the last
upon work done in the Society’s Prosectorium, dealt with the
rhinaria, the facial vibrisse, and the pads and interdigital integu-
ment of the feet in many of the genera of Canidez and all the
admitted genera of Urside. In connection with the Canide the
most interesting results were the discovery of evidence, supplied
by the feet, of kinship between Cuon and Lycaon, and between
Otocyon and Vulpes. On the other hand, two of the South
American fox-like dogs Cerdocyon microtis and Bendalopes: gracilis
differ widely from each other in the structure of the feet, and
equally widely from Vulpes and Otocyon. The most aberrant of
all the dogs in the matter of foot-structure is Speothos (Icticyon),
which is unique in having the digital pads of the third and
fourth toes of both front and hind feet united, and in having the
pollex of the front foot set low down adjacent to the postero-
internal angle of the main or plantar pad. These differences
seem to justify the division of the Canide into the two sub-
families Speothoine and Canine.
In the Urside four well-marked genera are characterised by
the structure of the feet, namely, Zhalarctos for maritimus, Ursus
for arctos and its allies horribilis and americanus, Tremarctos for
thibetanus, japonicus, ornatus, and malayanus, and Melursus tor
ursinus. Melursus differs from the rest in having all the digital
pads united almost to their distal ends. Tremarctos differs from
Ursus in haying the area behind the main or plantar pad of the
fore foot quite naked, instead of thickly hairy as in Ursus and
Thalarctos, and Thalarctos differs from Ursus in having the
plantar pads wide and short and the area of the sole behind the
plantar pad of the posterior foot covered with hair except for
a small lozenge-shaped pad towards its outer border. From
Tremarctos, malayanus may be eliminated as Helarctos by the
structure of the rhinarium. All the generic names adopted in
the paper had been previously proposed, mainly upon cranial and
dental characters; but the evidence from this source is so un-
satisfactory that there has been no agreement amongst zoologists
in their recognition.
Dr. G. A. Bounznerr, F.B.S., F.Z.8., contributed a paper
entitled “A Second Collection of Batrachians and Reptiles made
by Dr. H. G. F. Spurrell, F.Z.S., in the Choco, Colombia,”
48
A paper on Procolophon trigoniceps, a Cotylosaurian Reptile
from South Africa, was received from Mr. D. M. 8. Warson,
M.Se., F.Z.8.
Mr. A. W. WartzErs presented a paper on the “‘ Marine Fauna
of British East Africa and Zanzibar, from Collections made by
Cyril Crossland, M.A., B.Sc., F.Z.8., in the Years 1901-1902:
Bryozoa—Cyclostomata, Ctenostomata, and Endoprocta.”
Out of the twenty-four species from these three groups, four
are new; and, as the species mentioned are all from 10 fathoms
or under, it will not occasion surprise that the number of Cyclo-
stomata is but small.
Sections of the ovicells of Hntalophora and Jdmonea are figured,
and stress is again laid on the importance of the ovicells in the
classification of the Cyclostomata, and it is also thought that
the primary and early zocecia may give valuable help. The ovi-
cells and other points indicate that some “ Yntalophora” have
Diastoporidan characters necessitating separation. General par-
ticulars are given as to the ovicells of the Cyclostomata.
It is shown that the rosette-plate of Zoobotryon pellucidum,
described by Reichert as having a central pore with 8-9 pores
round it, has only one pore, but that surrounding it there are a
number of cells, so that there is a rosette of cells instead of
a rosette of pores. Also the groups of cells in the neighbourhood
of the rosette-plate have a fairly definite form, changing with the
species in some cases, thus furnishing specific characters.
In a note, further proof is brought forward showing that the
genus Lagenipora Hincks should be retained for a group recently
included in Cellepora, and Siniopelta Levinsen is a synonym.
Also the Celleporella Norman is Lagenipora,
This Meeting closes the Session 1913-1914, The next Meeting
of the Society for Scientific Business will be held on Tuesday,
October 27th, 1914, at half-past Frve o’clock p.m.
The following papers have been received :—
W. L. Distant.
eee
Report on the Rhynchota collected by the Wollaston Ex-
pedition in Dutch New Guinea.
49
T. H. Wiruers, F.G.S.
A remarkable new Cirripede from the Chalk of Surrey and
Hertfordshire.
F, A. Ports, M.A.
Polycheta from the N.E. Pacific: The Chetopteride. With
an Account of the Phenomenon of Asexual Reproduction in
Phyllochetopterus and the Description of Two new Species of
Chetopteride from the Atlantic.
F. EH. Bepparp, M.A., D.Sc., F.R.S., F.Z.8.
Contributions to the Anatomy and Systematic Arrangement
of the Cestoidea.—XV. On a new Genus and Species of the
Family Acoleide.
D. M.S. Watson, M.Sce., F.Z.S.
(1) A new Fossil Reptile from South Africa.
(2) Notes on some Carnivorous Therapsids.
(3) Hunotosaurus africanus Seeley, and the Ancestry of the
Chelonia.
E. Heron-Auten, F.L.S., F.G.8., F.R.M.S., and Arraur EHar-
LAND, F.R.M.S.
The Foraminifera of the Kerimba Archipelago, Portuguese
Kast Africa.
Communications intended for the Scientific Meetings should
be addressed to
P. CHALMERS MITCHELL,
Secretary.
ZOOLOGICAL Society oF Lonpon,
ReEGENtT’s Parn, Lonpon, N.W.
June 16th, 1914,
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No. 136.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.*
Octcber 27th, 1914.
Prof. E. A. Mixon, M.A., F.R.S., Vice-President,
in the Chair.
The Minutes of the last Scientific Meeting were confirmed.
The Secretary read a Report on the Additions to the Society’s
Menagerie during the months of June, July, August, and
September, 1914.
Mr. R. H. Burne, M.A., V.P.Z.S8., exhibited a number of
preparations showing some adaptations for the nourishinent of
the embryos of Hlasmobranchs,
Mr. R. E. Savacez exhibited two abnormal Herrings, taken by
trawl in the North Sea. The first specimen had neither pelvic
fins nor girdle (pelvic bones). ‘Phe usual position of base of fins
was indicated externally by the presence of the characteristic
elongated scales. The usual musculature was present internally.
‘The second specimen lacked the lett pelvic fin and pelvic bone ;
the musculature was complete.
Messrs. E. Heron-Auten, F.L.S., F.Z.S., and Arraur Har-
LAND, F.R.M.S., read a paper on the Horaminifera of the Kerimba
Archipelago, obtained by Dr. J. J. Simpson in the years 1907-8.
The area is a new one so far as the Foraminifera are concerned,
the on’y records in any way approximating to it being the species
described by d’Orbigny in 1826, by Brady in 1876 and 1884, by
Mobius in 1880, and by Egger in 1893, from material which was
collected from adjacent areas to the Hast of Madagascar, and otf
Mauritius and the Seychelles.
* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent's Park, N.W., on the Tuesday following the date of Meetung to which
it refers. It will be issued, along with the * Proceedings,’ tree of extra charge,
to all Fellows who subscribe to the Publications; but it may be obtained on the
day of publication at the price of Sixpence, or, if desired, sent post-free for
the sum of Sie Shillings per annum, payable in advance,
52
The material consisted of fine siftings from dredgings, and
having but few molluscan fragments and stones the larger
adherent forms are poorly represented, but 470 species and
varieties have been identified, including two new genera, and
28 new species and varieties. The general facies is strikingly
similar to that characteristic of Australian, Torres Straits, and
Malay gatherings. The problem of distribution thus raised is
obscure, the intervening ocean being abyssal, while the species
now recorded are all shallow-water types. Many of the special-
ized forms common to these widely separated areas do not
apparently occur in similar dredgings from intervening coasts
such as the Red and Arabian Seas. No doubt the Equatorial
Current, which traverses the Indian Ocean from E. to W.
and impinges on the African coast in our area, is primarily
responsible for this phenomenon.
Among the striking forms described and exhibited were
Ammodiscus charoides (Jones & Parker), now recorded for the
first time as a tropical species; Carterina spiculotesta (Carter),
exhibiting a hitherto unrecorded arrangement of the spicules ;
Pavonina flabelliformis WOrbigny, originally discovered by
d’Orbigny in sand from Madagascar in 1826 and lost sight of for
fifty years; Chrysalidina dimorpha Brady, a rare and beautiful
Textularian ; Discorbina dimidiata Parker & Jones, and Dis-
corbina polystomelloides Parker & Jones; Cymbalopora bulloides
1’ Orbigny, the dual nature of the terminal chamber, being divided
into a “balloon” and a contained ‘ float” chamber, was des-
cribed; also the species Cymbalopora millettii Heron-Allen &
Earland, first recorded from the Malay Archipelago by Mr. F. W.
Millett. A new record was established for Haddonia torresiensis
Chapman, hitherto only found in the Torres Straits and the
Tropical Pacific.
This paper will be published in the ‘ Transactions.’
Mr. T. H. Wirners, F.G.S., described a new Cirripede based
on a number of disconnected valves from the Chalk of Surrey
and a complete specimen from the Chalk of Hertfordshire.
Except for three valves referred to a new species of Scalpellum
(sens lato), the whole of the material belongs to a remarkable
new asymmetrical Cirripede which differs from Verrueca in the
more primitive structure of the valves, in the presence of two
lower lateral valves on the rostro-carinal side, and in the absence
of interlocking ribs. This species undoubtedly represents the
ancestral type from which has arisen the recent group of asym-
metrival sessile Cirripedes forming the family Verrucide, and in
its structure clearly shows its origin from the symmetrical
pedunculate Cirripedes of the family Pollicipedide. It presents
further evidence that the sessile condition was arrived at inde-
pendently on several different lines of descent during the evolution
of the Cirripedia.
53
Mr. W. lL. Disrant communicated his report on the Rhyn-
chota collected by the Wollaston Expedition in Dutch New
Guinea.
This paper will be published in the ‘ Transactions.’
The next Meeting of the Society for Scientific Business will
be held on Tuesday, November 10th, 1914, at half-past Five
o’clock P.m., when the following communications will be made :—
Lewis BALrour.
Exhibition of Photographs illustrating the Life-history of
the Garnet.
R. I. Pococg, F.RS., F.LS., F.Z.S.
Lantern exhibition showing the Differences between the
Pine and Beech Martens.
F. E. Bepparp, MEA: D.Se., F.R.S., E.Z.8.
Contributions to the Anatomy and Systematic Arrangement
of the Cestoide,.— XV. On a new Genus and Species of the
Family Acoleide.
Report on the Spiders collected by the British Ornitholo-
gists’ Union Expedition and the Wollaston Expedition to
Dutch New Guinea.
The following papers have been received :—
F. A. Porrs, M.A.
Polycheta from the N.E. Pacific: The Cheetopteride. With
an Account of the Phenomenon of Asexual Reproduction in
Phayllochetopterus and the Description of Two new Species of
Cheetopteride from the Atlantic.
D. M. S. Watson, M.Sce., F.Z.8:
(1) A new Fossil Reptile from South Africa.
(2) Notes on some Carnivorous Therapsids.
(3) Eunotosaurus africanus Seeley, and the Ancestry of the
Chelonia,
54
KATHLEEN HADDEN.
On the Methods of Feeding and the Mouth-parts of the
Larva of the Glow-worm (Lampyris noctiluca).
Gu Stuwanoson, Busy MD Eee
On Two new Subgenera of Freshwater Entomostraca.
Wituram Nicout, M.A., D.Sec., M.D.. F.Z.S.
A new Liver-fluke (Platynosomum acuminatum) from the
Kestrel.
Lt.-Col. J. M. Fawcerr.
Notes on a small Collection of Heterocera made by Mr. W.
Feather in British East Africa, 1911-12.
J. F. Gemuityi, M.A., D.Sce., F.Z.S.
Abnormal Gills in the Starfish, Porania pulvillus O. F. M.
F. F. Larpiaw, M.A., F.Z.S.
Contributions to a Study of the Dragonfly Fauna of Borneo.
Part III. A Collection made on Mount Kina Balu by
My. Moulton in September and October 1913.
Communications intended for the Scientific Meetings should
be addressed to
P. CHALMERS MITCHELL,
Secretary.
ZOOLOGICAL Society or Lonpon,
Recenv’s Park, Lonpon, N.W.
November 3rd, 1914.
No. 1387.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.”
November 10th, 1914.
Prof. E. W. MacBripve, M.A., D.Sc., F.R.S., Vice-President,
in the Chair.
The Minutes of the last Scientific Meeting were confirmed.
The Secretary read a Report on the Additions to the Society’s
Menagerie during the month of October.
The SrcreTary exhibited a photograph showing Oysters
eee upon mangroves at Lobito Bay, Portuguese West
rica.
Mr. L. H. Jaugs, F.Z.S., communicated some notes upon the
birth of a Porpoise at the Brighton Aquarium.
Mr. R. I. Pocock, F.R.S., F.L.S., Curator of Mammals, gave
an exhibition, illustrated by lantern-slides, showing some un-
recorded structural differences between the Pine-Marten (J/artes
martes) and the Beech-Marten (artes foina), and pointed out
that the two species, apart from the known differences in the
skull and teeth, may be distinguished by the size of the ears,
which are broader and longer in J. martes than in JZ. foina,
and by the dimensions of the pads on the feet, which are con-
siderably larger and less overgrown with hair in J/, foina than
in WM. martes.
* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent’s Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications ; but it may be obtained on the
day of publication at the price of Strpence, or, if desired, sent post-free for
the sum of Six Shillings per annum, payable in advance,
56
Mr. R. H. Burne, M.A., V.P.Z.S., exhibited some paraffin
Simulacra of Molluscan and other shells made accidentally by
Dr. G. V. Ariens Kappers while embedding objects for the
microtome. During this process, paraffin in a molten state
accidentally escaped from the mould and set in the shell-forms
shown, probably owing to distorted crystallization.
Dr. F. E. Bepparp, M.A., F.R.S., Prosector to the Society,
read a paper on the Anatomy and Systematic Arrangement of
the Cestoidea, in which he described a new genus and species
of the Family Acoleide, based upon a large number of examples
obtained from the Canadian Tree-Porcupine (Hrethizon dor-
satum).
Mr. H. R. Hoae, M.A., F.Z.8., read his report on the Spiders
collected by the Wollaston and British Ornithological Union
Expeditions in Dutch New Guinea. This collection confirms a
good deal of the work of previous authors, and at the same time
brings to light much that is new. Leaving out the Attide, there
are representatives of nine families, comprising twenty-six genera
(of which one is new) and forty-five species or subspecies of which
some nineteen are new. Included among the latter are the
following :—
CoNOTHELE SPINOSA, Sp. n.
Differs from C. malayana Dol. in having the front lateral eyes twice as long as the
rear laterals, the rear median smaller still, and the front median 14 times as broad
as the rear, instead of all the eyes equal. From C. arboricola Poc. in having the
first pair of legs longest instead of the fourth, and four teeth only on the outer
margin of the falx-sheath instead of six. From C. doleschalli Thor. similarly in
the legs, and the latter has the rear median and side eyes close together.
SELENOCOSMIA LANCEOLATA, sp. n."
Differs from S. similis Kule. and A. honesta Hirst, possibly the same, in having
the patella cum tibia and metatarsus cum tarsus about the same length as, or
shorter than, the femur and trochanter in the first three pairs of legs instead of much
longer. The abdomen is paler and the lip and maxille the same colour all over
instead of being particoloured.
Length 34 mm.
PsECHRUS CASTANEUS, Sp. 0.
Differs from P, argentatus Thor., P. libeltii Kule., and P. annulatus Kule. in
having the legs longer in proportion, the front pair being more than ten times as
long as the cephalothorax instead of about eight, in having the front median eyes
as large as the side instead of smaller, and the rear median eyes more than their
diameter apart instead of less. It is also only two-thirds the size of the first-named
and larger than the last.
Length 17 mm.
FPECENIA CINEREA, sp. n.
Differs from F’. swmatrana Kale. in its first pair of legs eight times the length of
the cephalothorax instead of about six, in the median eye-square broader than long,
and the rear row distinctly procurved instead of nearly straight. The epigyne also
differs considerably, though rather near to Mr. Rainbow’s drawing of his F. oblonga
from the Solomon Islands.
Length 10 mm.
57
ARANEUS FLORIATUS, sp. n,
Differs from A. pfeifferi Thor. and JA. ferruginea Thor. (‘ Ragni di Selebes,
pp. 35, 38), besides a more elaborate back pattern, in having the median eye-area
broader than long instead of longer than broad, and the median eyes distant from
the side eyes by twice instead of three times their distance between one another,
also scape of epigyne four times as long as its greatest breadth instead of twice.
2 females, 1 male. Length 17 mm.
ARANEUS GRANTI, sp. n.
Differs from A. vatius Thor. (loc. cit. supra, p. 41), which it rather closely
resembles, in having the front median eyes their diameter apart instead of much
more. The sternum is bright cream-colour instead of dark yellow-brown. The
scape of the epigyne issues from the upper part of the corpus instead of the lower
margin, is 23 times as long as the width of the corpus instead of twice, and in the
middle widens to twice its narrowest part above instead of being straight, four times
instead of twice as long as broad.
1 female. Length 125 mm.
LEUCAUGE CAUDATA, Sp. n.
Differs from Araneus caudifer Kule., which it rather resembles, in its smaller
size, the,legs only half as long, its colouring silvery imstead of reddish brown, the
year median eyes half their diameter apart instead of more than their diameter, so
that the area of the median eyes is narrowest instead of widest posteriorly. ‘The
epigyne has a bulbous base, wanting in the other, but a shorter scape.
Length 8 mm.
REGILLUS DIVERGENS, sp. n.
Differs from R. asper Camb. in its larger size, the eyes of the rear row slightly
recurved instead of straight, equidistant instead of median nearer together than they
are from the side, thus forming with the front median eyes a square instead of
trapezium narrowest posteriorly.
Length 12 mm.
OLIOS PRINCEPS, sp. n.
Cephalothorax chestnut-red with pale red-brown bristles and hair. Abdomen
pale yellow-brown with faint median and branching stripes of a darker colour, short
longitudinal flecks of the same on the sides merging into longer underneath to form
a shield-shaped pattern, above and just below the genital fold dark brown, on the
latter a streak of white. Mandibles, lip, and maxille black-brown; sternum bright
red, legs and palpi bright red-brown with brown hairs, that on the coxe black-
brown. Vulva a broad chitinous horseshoe-frame, widely open at the base, the
hollowed inner area indistinctly divided by a median longitudinal fovea.
1 female. Length 23 mm.
OLIOS ACTHON, sp. n.
Differs from Olios salacius L. K. in having the median eyes of the rear row
(which is slightly procurved) distinctly farther apart than they are from the front
median. The sternum bright yellow with eight black hair-spots ranged round just
inside the margin instead of brownish yellow with white hair-stripes, pattern on
underside of abdomen like Olios (Het.) diana L. K. Five large teeth and one
smaller on the inner falx-margin and two on the outer, instead of three and one
respectively. The palp is furnished with a spiral of ten turns round the usual drum.
It is also rather smaller.
1lmale. Length 133 mm.
PALYSTES DASYURINUS, sp. n.
Differs from P. ignicomus L. K. in having ten white hair-spots on black shield
on underside of the abdomen, front median eyes their diameter apart and half that
distance from the laterals, three large teeth and one rather smaller on inner margin
of falx-sheath, one large between two smaller on outer—a single, long, curved bristle
on upper inner margin of falx.
lfemale. Length 25 mm.
HETEROPODA VENATORIA Linn., var. PLURIDENTATA, Noy.
Differs from usual form in having five large teeth and one smaller on inner
margin of falx-sheath, one medium-sized and two smaller on outer margin.
2 females.
58
EXOPALYSTES, gen. nov.
Intermediate between the groups Delenee and Heteropodew, but near Palystes
L. K., differing therefrom in having the front median eyes as large as the side eyes,
the eyes of the front row larger than those of the rear row, the eyes all sessile and
the clypeus about twice as broad as a front median eye. The cephalothorax, highest
in the posterior one-third, sloping to the front, and a thick fringe of long bristles on
the upper inner margin of falx instead of one single bristle.
EXoPALYSTES PULCHELLA, sp. n.
Pale yellow with white hair all over, except underside of femora i. and ii, which
is dark grey. Here and under the tibia of the same pairs a thick mat of long
yvecumbent cylindrical bristles on the anterior two-thirds of the joints.
1 female. Length 18 mm.
The next Meeting of the Society for Scientific Business will
be held on Tuesday, November 24th, 1914, at half-past Five
o'clock P.m., when the following communications will be made :—
EXHIBITIONS AND NOTICES.
EK. Heron-Axen, F.L.S., F.Z.8., and Arruur HARLAND, F.R.M.S.
Exhibition of Tests of Arenaceous Foraminifera to introduce
a Discussion on the Interpretation of these Structures.
D. M.S. Watson, M.S8ce., F.Z.S.
(1) A new Fossil Reptile from South Africa.
(2) Notes on some Carnivorous Therapsids.
(3) Hunotosaurus africanus Seeley, and the Ancestry of the
Chelonia.
A, Ports, M.A.
Polycheta from the N.E. Pacific: The Chetopteride. With
an Account of the Phenomenon of Asexual! Reproduction in
Phyllochetopterus and the Description of Two new Species of ©
Cheetopteride from the Atlantic. ;
59
The following papers have been received :—
KATHLEEN HADDEN.
On the Methods of Feeding and the Mouth-parts of the
Larva of the Glow-worm (Lampyris noctiluca).
G. Srewarpson Brapy, M.D., LL.D., F.R.S., C.M.Z.8.
On Two new Subgenera of Freshwater Entomostraca.
Wattram Nicout, M.A., D.Sc. M-D., F.ZS.
A new Liver-Fluke from the Kestrel.
Lt.-Col. J. M. Fawcett.
Notes on a small Collection of Heterocera made by Mr. W.
Feather in British East Africa, 1911-12.
J. F, Gemuiut, M.A., D.Sc., F.Z.8.
—————EE ane
(1) Abnormal Gills in the Starfish, Porania pulvillus O. F. M.
(2) On the Ciliation of Asterids, and on the Question of
Ciliary Nutrition in Certain Species.
F. F. Larpiaw, M.A., F.Z.8.
ss =
Contributions to a Study of the Dragonfly Fauna of Borneo.
—Part ITI. A Collection made on Mount Kina Balu by
Mr. Moulton in September and October 1913.
E. G. Boutencer, F.Z.S.
On a Colubrid Snake (Xenodon) with a vertically movable
Maxillary Bone.
Communications intended for the Scientific Meetings should
be addressed to
P. CHALMERS MITCHELL,
Secretary.
ZooLoGIcAL Society oF Lonpon,
Recent’s ParK, Lonpon, N.W.
November 17th, 1914.
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No. 188.
ABSTRACT OF THE PROCEEDINGS
ZOOLOGICAL SOCIETY OF LONDON.*
November 24th, 1914.
Prof. E. A. Mincury, M.A., F.R.S., Vice-President, in the Chair.
The Minutes of the last Scientific Meeting were confirmed.
Dr. R. Broom, C.M.Z.S., exhibited the skull of a new type of
Thecodont Reptile from the Upper Permian Beds of South Africa,
and a number of skulls of Zrichosurus vulpecula, Phascolarctus
cinereus, Chrysochloris hottentota, and C’. asiatica, illustrating
dental variations.
Mr. D. Seru-Smitu, F.Z.S., Curator of Birds, exhibited an egg
of the New Guinea Rifle-bird (Ptilorhis intercedens), which had
been laid in the Society’s Gardens in July last, the first instance
of any species of Paradise-Bird laying in the Gardens.
Mr. E. T. Newron, F.R.S., F.Z.S., exhibited a series of bones
of animals showing indications of natural repair, and a number
of teeth of a female Sperm- Whale (Physeter macrocephalus).
Dr. C. W. Anprews, F.R.S., F.Z.8., gave an account of three
papers by Mr. D. M.S. Watson.
The first paper contained the description of a new reptile from
the Permian of the Cape Province, 8. Africa, which Mr. Watson
regards as derived from a Cotylosaurian ancestor and as perhaps
related to Areoscelis and the modern lizards. A new genus is
* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent's Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications ; but it may be obtained on the
day of publication at the price of Sixpence, or, if desired, sent post-free for
the sum of Stix Shillings per annum, payable in advance.
62
founded for the reception of the so-called ‘ Proterosaurus
hualeyi.”
In the second paper the origin of the Chelonia is discussed,
and a number of reasons given for supposing that they may be
descended from some such form as Hunotosaurus africanus
Seeley.
In the third paper Mr. Watson describes the skulls of Bauria,
Microgomphodon, and Sesamodon, and discusses the relation of
the group with the Cynognathids. He also describes a new
skull of Lycosuchus, in which both the prevomers and vomer are
present.
Mr. F. A. Porrs, M.A., communicated a paper entitled
“ Polycheta from the N.E. Pacific: The Chetopteride. With
an Account of the Phenomenon of Asexual Reproduction in
Phyllochetopterus and the Description of Two new Species of
Chetopteride from the Atlantic.”
The new species of Phyllochetopterus was found in branched
tubes, each usually containing several individuals. The origin of
these colonies each from a single individual is suggested by the
frequent occurrence of worms in various stages of regeneration.
An examination of these shows that autotomy first occurs in the
middle region of the animal’s body, and a complete animal is
regenerated from each of the two parts. ‘This phenomenon
appears to be characteristic also of another new species of this
genus from Plymouth, which lives in small colonies in branched
tubes.
Several points in the morphology of the Cheetopteride are also
discussed.
Messrs. E. Heron-Axten, F.L.S., F.Z.S., and Artaur HARLAND,
F.R.M.S., exhibited a series of microscopic preparations and
photographic views of the tests of Arenaceous Foraminifera, and
utged their view that these afforded evidence of purpose and
intelligence on the part of the Foraminifera.
An interesting discussion followed, in which, amongst others,
Sir H. H. Howorth, F.R.S., Sir E. Ray Lankester, F.R.S., and
the Secretary took part.
63
The next Meeting of the Society for Scientific Business will
be held on Tuesday, February 9th, 1915, at half-past Five
o’clock p.m. ‘The Agenda for this Meeting will be circulated early
in January.
The following papers have been received :—
KATHLEEN HADDON.
On the Methods of Feeding and the Mouth-parts of the
Larva of the Glow-worm (Lampyris noctiluca).
G. _STEWARDSON Brapy, M. D., LL. iDY5 1 R. Se C. M. ZS.
On T ‘wo new Subgenera, ie Dives iawrnere ison Ae.
A new Liver-Fluke from the Kestrel.
Lt.-Col. J. M. Fawcerrt.
Notes on a small Collection of Heterocera made by Mr. W.
Feather in British Hast Africa, 1911-12.
(1) Abnormal Gills in the Starfish, Porania pulvillus O. F. M.
(2) On the Ciliation of Asterids, and on the Question of
Ciliary Nutrition in Certain Species.
F. F. Larpriaw, M.A., F.Z.8.
Contributions to a Study of the Dragonfly Fauna of Borneo.
—Part II. A Collection made on Mount Kina Balu by
Mr. Moulton in September and October 1913.
E. G. BouLencsr, F.Z.8.
On a Colubrid Snake (XYenodon) with a vertically movable
Maxillary Bone.
RowtAnn H. TurNER.
Descriptions of New Fossorial Wasps from Australia,
Communications intended for the Ss
be addressed to oat
P, CHALMERS MITC
- *
ZOOLOGICAL Society or Lonpon, °
Recent’s Park, Lonpon, N.W.
December 1st, 1914,
38.
39.
40.
41.
42.
43.
44,
45.
Papers (continued),
Cephalopoda from the Monte Bello Islands. By G.O. Rossoy, B.A. (Text-figure 1.)
Description of a new Lizard from the Canary Islands. By Dr. Pu. Lmurs. .........
The Mechanism of Suction in the Potato Capsid Bug, Lygus pabulinus Linn. By
P. R. Awarr, B.A. (Cantab.), D.I.C. (Lond.), Sir John Wolfe-Barry Research Scholar,
Imperial College of Science, London. (Text-figures 1-29.)
Se ere Pee ese sen teen eese
Procolophon trigoniceps, a Cotylosaurian Reptile from South Africa. By D. M.S.
Watson, M.Sce., F.Z.S., Lecturer on Vertebrate Paleontology in University College,
ondon. (Plates ILE and Poxt-mgures tO.) isis. va veins esgic pelecieecoss ches
The Deinocephalia, an Order of Mammal-like Reptiles. By D. M.S. Watson, M.Sc.,
F.Z.8., Lecturer on Vertebrate Palzeontology in University College, London. (Plates
Vee AG neat ferns al Gale ehare aie tater oye da sia Wo era's vlelerel ei auve.a © a7 a Waln i tie sare oom
Diagnoses of New Genera and Species of Zonitide from Equatorial Africa. By H. B.
RESTO Nes HZ Satan (Cll vie syle bes) Rar aratetrarehcr ccrcticveke't;ercveisie|e, cree ou wis tei a'e ateiehelani ciawiwieroe
On a second Collection of Batrachians and Reptiles made by Dr. H. G. F. Spurrell,
F.Z.8., in the Choco, Colombia. By G. A. Bounznenur, F.R.S., F.Z.8. (Plates
PECL T Seo crsucretercrersteleie ees ays Brere tr eE Mere agate Sr ate ots (eee OU or ctasola ate, 5 Pe etal ei avere: wsilac ets
Zoological Results of the Third Tanganyika Expedition, conducted by Dr. W. A.
Cunnington, 1904-1905.—Report on.the Parasitic Eucopepoda. By Wruiam A.
' Cunnineton, M.A., Ph.D., F.Z.8. (Plate I., and Text-figure 1.) .......... 0.0000
46.
Ae
48,
49.
The Marine Fauna of British East Africa and Zanzibar, from Collections made by
Cyril Crossland, M.A., B.Sce., F.Z.S., in the Years 1901-1902. Bryozoa—Cyclo-
stomata, Otenostomata, and Endoprocta. By Arruur Wm. Waters, F.L.S., F.G.S,
(latest Vian dallont streaney Uy \eeinitcrerars/stefsicle eisl vies s| olelelel cove sjale«'ociehels 2/3 etalevale oe
Contributions to the Anatomy and Systematic Arrangement of the Cestoidea.—
XIV. On a new Species of Rhabdometra, and on the Paruterine organ in Otiditenia.
By Frank H. Bupparp, M.A., D.Sc., F.R.S., F.Z.S., Prosector to the Society. (Text-
MOTOS USM icneopeconicsobo ey do deigs ome OS TIDO BOUND DOOUOE a OG uEmEmn uD contion oc
On the Facial Vibrisse of Mammalia. By R. I. Pocock, F.RS., F.LS., F.Z.S.,
@urator of Mammals (Rex nemrestl—19.)) sce c ed eee eet ev cv dbeee ee adanne
On the Feet and other External Features of the Canidz and Urside. By R. 1. Pococr,
F.R.S., F.L.S., F.Z.8., Curator of Mammals (Text-figures 1-13.) ..........+- coee
Page
677
681
685
813
819
831
859°
889
pds}
jot
)
LIST OF PLATES,
1914, Parr III. (pp. 491-94).
a
H IXLEY <= Pl. 7
Seah II } Courting-habits of Podiceps cristatus .....
Parkins : Ply el: Structural characters of Paralastor Sat
Monraaun: Phe
Ratipun : Pl. | Crustacea from the Monte Bello Islands .
Watson: ‘nd Ope f ors: sé
II. | Procolophon trigonicepS ....+4.eeeereee:
iBHE, Fae
Warson: ded lage AM
WV:
PRESTON: PL ees ade
Bounencer: _ Pl. I. 1. Atelopus spurrelli, 2, inptccblophane
intermedius. 3. Polychrus spurrelli ...
II. 1. Leptophis brevior. 2. Homalocrani
Cunsincron: Pl 1. 1-8. Lerneocera diceracephala, 4-7. L. ha
plocephala. 8,9. L. temnocephala...
Waters: I; ° :
ah aoe from Zanzibar......00..5.
ay:
NOTICE.
so that the es reference i is now P. Z. 8. 1914, Dp. " fei
is as follows :— Ln af
~ Part Tsasenedl in March.
ty he See ys Ons! | : fash
» III. ,, September, —
TV. -
»
is dt cea
a
PROCEEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF THE
‘ZOOLOGICAL SOCIETY
OF LONDON.
1914.
PART IV:
containine Paces 945 to 1077, witH 9 PLATES,
36 Tuxt-ricuREs, TiITLEPAGE, INDEX, ETC.
oy
DECEMBER (i FEB 25 1915
PRINTED FOR THE SOCIETY,
SOLD AT uae HOUSE IN REGENT’S PARK.
LONDON :
MESSRS. eee NS, GREEN, AND CO.,
PATERNOSTER ROW.
Slay ab |
Sse mN
Se) a !
[Price Twelve Shillings. | aes
= aes = p20
LIST: OF CONTESTS
1914, Part LV. (pp. 945-1077).
EXHIBITIONS AND NOTICES.
Page
The Secrerary. Report on the Additions to the Society's Menagerie during the months
of June, July, August, and September, 1914 ...--.-e.see cece cece cence ee eeeees 1057
Mr. R. H. Burne, M.A., V.P.Z.S. Exhibition of some adaptations for the nourishment of
embryos of Elasmobranchs 2... ..s0scnercsoeecs cere eeeens ee stseucss oeumee oniue 1059
Mr. R. E. Savace, Exhibition of two abnormal Herrings .....-....--..-+...e+eeeees 1060
Messrs. E. Heroy-Auten, F.LS., F.Z.S., and Artuur Earuanp, F.R.M.S, Notice of
Report on the Foraminifera of the Kerimba Archipelago, obtained by Dr. J. J.
Sinmipson in the years 1907-6 2... nce se ccieg se ec awiseist sivas ng sy ae nee ee ++» 1060
Mr. W. L. Distanr. Notice of Report on the Rhynchota collected by the Wollaston
Expedition in Dutch New Guinea ............-+ sees ona weed sia ses 5: get ate coe
The Sscrerary, Report on the Additions to the Society's Menagerie during the month
Of Oebober VOLS 2 o.2 scine sieistarcin saln's Silos ono aalisle es lwinigip ate epee eeeees 1060
The Secretary. Exhibition of a photograph of Oysters growing upon mangroves at
Hobito. Bay, Portuguese" Weestiatriea:jctec <> .0l< s-1aiaiaieleiev seein jo earn Seo) ole ee ana 1061
Mr. R. H. Burne, M.A., V.P.Z.S. Exhibition of Simulacra of Molluscan Shells ....... » L061
Mr. H. R. Hoge, M.A., F.Z.8. Notice of Report on the Spiders collected by the British
Ornithologists’ Union and Wollaston Expeditions in Dutch New Guinea .......... 1061
Mr. Lewis H. James, B.A.,F.Z.8. Notes on the birth of a Porpoise at Brighton Aquarium. 1061
Mr. R. I. Pococs, F.R.S., F.L.S., F.Z.8., Curator of Mammals. Exhibition showing new
points of difference between Pine and Beech Martens, (Text-figures 1-4.) ........ 1062
Mr. E. T. Nzwron, F.R.S., F.Z.S. Exhibition of bones of animals showing indications of
natural repair, and some teeth of a female Physeter ...........0.0000es ‘sietendeneeanes . 1069
Messrs. E. Heroy-Aurey, F.L.S., F.Z.S., and Arruur Earuanp, F.R.M.S. Exhibitionand _
Discussion on “ Purpose” and “Intelligence” in the Foraminifera ..............-- 1069.
Contents continued on page 3 of Wrapper. —
ZOOLOGICAL SOCIETY OF LONDON:
Turs Society was founded in 1826 by Sir Sramrorp RaFFLEs,
Mr. J. Sasrve, Mr. N. A. Vicors, and other eminent Naturalists,
for the advancement of Zoology and Animal Physiology, and for the
introduction of new and curious subjects of the Animal Kingdom,
and was incorporated by Royal Charter in 1829.
instit
v t lg
Dy
e
Patron.
HIS MAJESTY THE KING.
COUNCIL.
HIS GRACE THE DUKE OF BEDFORD, K.G., F.R.S., President.
Ricuarp H. Burne, Esa., M.A.,
Vice-President.
Aurrep H. Cocks, Ese., M.A.
Tor Rr. Hon. Tor Earn or
Cromer, EC. -G.C.B.,
G.C.M.G., K.C.S.1., F.R:S.,
E. G. B. Mrapr-Watpo, Esa.
Prormssor Epwarp A. Minceiy,
M.A., F.R.S., Vice-President. j
P. Costmers Mrrenent, Ese.,
MAL, D.Sc. LL:D., FRS.;
Vice-President.
F. G. Dawrrey Drewrtt, Esa.,
M.A., M.D.
Crartes Drummonn, Kse.,
Treasurer.
Turn Eart oF Dunmors, V.C.,
M.V.O.
Sir WarteR Roper Lawrence,
Br., G.C.LE., Vice-President.
Ervest W. MacBrinz, Esa.,
M.A., D.Sc., F.R.S., Vace-
President,
Secretary.
W. RB. Oeitvir-Grant, Ese.
Aubert Pam, Esa.
Appian D. W. Pottocg, Ese.
Tur Eart oF PorrsmoutTH.
Tur Maravess or Srico, F.S.A.
Oxtprietp Tuomas, Esa., F.R.S.
Antuony H. WrnGFIzLD, Hse.
Henry Woopwarp, Kse., LL.D.
F.R.S., Vice-President.
9
a
The Society consists of Fellows, and Honorary, Foreign, and
Corresponding Members, elected according to the By-Laws. It
carries out the objects of its foundation by means of the collection
of living animals, by its Library, and by its Scientific Publications.
The Office of the Society, Regent’s Park, N.W., where all com-
munications should be sent, addressed to ‘* The Secretary,” is open
from Ten till Five, except on Saturdays, when it closes at One p.m.
The Library, under the superintendence of Mr. Henry G. J. Peavot,
is open daily (except Sunday) from Ten a.m. till Five p.m.; on
Saturdays, Ten a.m. till Two p.m.
‘he Library is closed from Good Friday to Easter Monday, and
upon all other Bank Holidays. It is also closed annually for
cleaning purposes during the whole month of September.
The Meetings of the Society for General Business are held in the
Meeting Room at the Society’s Office on the third Wednesday in
every month of the year, except in September and Octeber, at half-
past Four o’clock p.m.
The Meetings for Scientific Business are held in the Meeting
Room at the Society’s Office fortnightly on Tuesdays, except in
July, August, September, and December and January, at half-past
Five o’clock p.m.
The Anniversary Meeting is held on the 29th. of April, or the
nearest convenient day, at Four p.m.
The Society’s Gardens are open daily from Nine o’clock until
Sunset. Mr. R. I. Pocock, F.R.S., F.LS., is the resident Super-
intendent and Curator of Mammals, Mr. D. Seth-Smith is Curator
of Birds and Inspector of Works, Mr. E. G. Boulenger is Curator
of Reptiles, and Prof. H. M. Lefroy is Curator of Insects.
The Prosectorium for Anatomical and Pathological work is under
the charge of Mr. Frank E. Beddard, M.A., D.&c., F.R.S., Prosector,
assisted by Mr. H. G. Piimmer, F.R.S., M.R.C.S., Pathologist to
the Society.
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ey)
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Secretary.
Regent’s Park, London, N.W.
December, 1914.
MEETINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON
FOR
SCIENTIFIC BUSINE Ss:
1915.
TuESDAY, FEBRUARY ...... 9 and 238.
Bye ENUM Hin ons ca) sceuc,0°2 9 4, -2or
= APRIL ed mene
x WON SRE Oe ee ea Diss, Deve
af RUNGE Ree res. 8
The Chair will be taken at half-past Five o’clock precisely.
AVOEOG IOC SOCIETY OF LONDON.
LIST OF PUBLICATIONS.
Tue scientific publications of the Zoological Society of London
are of two kinds—“ Proceedings,” published in an octavo
form, and ‘‘ Transactions,” in quarto.
According to the present arrangements, the “ Proceedings”’
contain not only notices of all business transacted at the scien-
tific meetings, but also all the papers read at such meetings
and recommended to be published in the “‘ Proceedings” by
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“ Proceedings,” to illustrate the new or otherwise remark-
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illustrations, figures of the new or rare species acquired in a
living state for the Society's Gardens are often given.
The “ Proceedings” for each year are issued in four parts,
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The following is a complete list of the publications of the
Society already issued.
TRANSACTIONS OF THE ZOOLOGICAL SOCIETY OF LONDON
4to. 19 vols. and Index. Price to Price to the
Fellows, Public.
Vols. I-IV. (out of print).
Vol. V., containing 67 Plates.. (1862-66) .... 5 4 6 19
3 VL, 4 2 yee a USCD=G0), Cor ealelaeees 15 0
Pee Liles + 73 sp, we LBC9= 72) 2 ame 13 12
be VALLE, rs 82 gp tae (AB72-74) 9 ae eS 12 Il
” Sy ” 99 29 (18/95-77) 12 1 16 2
i c i 95 39+, oe AISAT—70) ee LOO . Loge
Index, Vole (2K. Wa aoe aa (1833-79) .... at : 10
Vol. XI., containing 97 Plates .. (1880-85) .... 12 . 12 16
iin? 4 BEE 5 65 se as (AEBO=DOW Face 8 4
she OAR BES _ 62 yl es (Leo Oo) aries 8 im
et GN % 47 pe SO6- Ue) ie mer 5 0
5 Meee, ¥ 52 Paar Rett fom 1901) a 15 14
KWOOCOK NONI ®D oo ©
DOWWWADSDSOHOCODAGOHOWOCOAwWAWoow
HoHwanntaaMHeNwc
CO OO He
Soo sSoooocesooococecoeceoseococooc ]
Seo, ia, ieee ame oe OOO T= ana 8
PPP NeN sles cs Al 5, «« (1908-1906) % > 18 5 18
a> Gg 8 i) Pe mie) ») ae (C1907 19 eee J :
wad. an 24 i ee (1909-19 LON a . 13 12
sie ek — Dart, (Pls: T—-V.).. (eb: Lota yee 18 4
» 2 (Pls: VI-XV.) (April 1912)... 5 0
8. (Pls XV LX LL. Beh, 191: 3). 2 10
Vf A Pls) XX DV XX VIL ae 9 12
» o (Pls. XX VILT-XXX.) y(Mazohi914) 9 12
,, 6. (Pl. XXXI.) (March 1914) .. 3 Pe 5
a (eal ee. @.8o.41 1s XXXII )(May1914) 7 10
Fy veda, Wary LOA) od io aes) win eee ene 2 3
55 pn aONaa TOTA Rn a ar 2 3
Flos (Sune WOU) 345s eee 2 3
= dite (Pl RX) (Nov: 190A) fi 10
», 12. (Pls. XXXV.-XXXVII_.)(Nov.1914)9 12
In consequence of a re-arrangement of the stock of the ‘Transactions,’ the Society is
#now able to offer for sale, at the reduced price of £50, sets of Vols. V—XVE. inclusive, and
A eoanate papers, of which a list can be supplied, at. about one-fourth their published price.
PROCEEDINGS OF THE COMMITTEE OF SCIENCE AND
CORRESPONDENCE OF THE ZOOLOGICAL SOCIETY OF
LONDON. 8vo. 2 vols. (Letterpress only), Price te Price tor
Fellows. Public.
Part I. 1830-31. 1 vol. 8vo., out of print.
jell eae PP he oes teatitn ohctes eoaat As. (G0. isn tate
PROCEEDINGS OF THE ZOOLOGICAL SOCIETY OF LONDON.
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SEcoNnD SuExiEs.
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PZ. o. A914) WLS Se ee
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LSS ISSLOWAISIRON Wain eive.
10-12. SCALPELLUM VIMINEUM.
ON A REMARKABLE NEW CIRRIPEDE. 945
PAPERS.
50. A remarkable new Cirripede from the Chalk of Surre
I Ng
and Hertfordshire. By THomas H. Wiruers, F.G.S.*
[Received May 27, 1914: Read October 27, 1914.]
(Plate I.¢ and Text-figure 1.)
INDEX. Page -
DnGrocu cho nme een eer ct en Meee dA oe ed eee apk OAS
SUIPDKCH TOURS ON? JEVAOAA POMC cen aceaseensinenees eobbadesd seebacacaneel i a Fal0
Evolution of the Verrucide ............. 951
Distribution (Geological), Cretaceous at Tivelens ere
verruca, Scalpellum (sensu lato) .....0.........00.2..... 946, 952
Proverruca vineulum, gen. et sp. 0. ......... 946
Proverruca compared with Verruca fail. Calantivn
(Scaliiclieras) lteter eA oleae: 950)
Seal pellumwiminreumer SP. Ws... 122 sectecce see eeseee-te--s.au--- 902
Among some Cirripede remains recently collected from the
Chalk of | Surrey are a number of isolated valves, which, together
with a remarkably complete specimen of the same species from
the Chalk of Watford, Hertfordshire, throw much light on the
evolution of the sessile Cirripedes of the family Verrucide. They
constitute, in fact, the ‘“ missing link” between the pedunculate
Cirripedes of the family Pollicipedide and the sessile asym-
metrical Cirripedes of the family Verrucide.
The valves from Surrey were obtained by me from a mass of
unusually soft Chalk found by Mr. C. P. Chatwin in the Slines
Oak Pit, Worms Heath, near Woldingham, and to judge from the
single example of Alicraster precursor collected from the same
horizon, the chalk appears to be in the upper part of the zone
of Micraster cor-testudinarium. This is the zone to which it has
‘been assigned by Mr. G. W. Young f.
Much difficulty was experienced in studying these valves, for
although they were all somewhat similar in external ornament,
the scuta and terga differed so much in structure that it was
apparent that more than one species was represented. What was
evident, however, was that certain of the scuta and terga, appar-
ently of the same species, were extremely convex tr. ansversely and
formed a semicircle when placed in position. This fact suggested
to me the possibility of their belonging to a new form of sessile
Cirripede, since if the capitulum were completed it would approach
more closely to radial symmetry than is the case in an ordinary
* Communicated by Dr. W. T. Carman, F.Z.S.
+ For explanation of the Plate see p. 953.
~ 1905. “Phe Chalk Area of North-east Surrey,’ Proc. Geol. Assoc. London,
vol. xix. p. 208 (pit 127a).
Proc. Zoou, Soc.—i914, No. LXIV. 64
946 MR. T. H. WITHERS ON
pedunculate Cirripede. It was at this stage of my observations
that I went to examine some Cirripedes at the Museum of Prac-
tical Geology, and among them noticed a beautifully complete
example of the species to which most of the isolated valves belong.
This fine specimen was collected by Mr. J. Rhodes from the
“ Upper Chalk of Watford Tunnel,” * and was apparently obtained
from a hollow flint at about the same horizon as the isolated
valves from Surrey.
Through the kindness of the Director of the Geological Survey
and Dr. F. L. Kitchin, I was enabled to borrow this specimen for
description.
Although this new Cirripede occupies a position intermediate
between the Pollicipedide and the Verrucide, I do not think it
advisable to found a new family for its reception. It will, I
think, be sufficiently distinguished if it is placed in a new genus,
and the diagnosis of the family Verrucide extended to include it,
since, it is undoubtedly a primitive Verrucid.
The remaining valves, which obviously do not belong to this
Verrucid, are described as a new species of Scalpellum (sensu
lato).
Family VERRUCID# emend.
Sessile, asymmetrical, box-like* barnacles, in which a scutum,
tergum, rostrum, and carina, with or without a rostral- and a
cavinal-latus in addition, are immovably united to form the
“wall”; the remaining scutum and tergum are movable, and form
the lid-like top.
PROVERRUCA, gen. nov.
Verrucids in which a rostral- and a carinal-latus are present on
the rostro-carinal side, and in which none of the valves has
developed interlocking ribs.
PROVERRUCA VINCULUMYT, sp. n. (PI. I. figs. 1-9; Text-
fig. 1, C-F.)
Material.—(1) A single complete individual in which the moy-
able seutum is somewhat displaced, and the upper portion of the
rostral-latus broken away. (2) A number of isolated valves
comprising :—1 fixed scutum (left), 6 fixed terga (5 right and
1 left), 4 movable scuta (1 right and 3 left), 2 movable terga
(left), 1 carinal-latus (right), and 2 rostral lateral valves (left).
There are no isolated examples of the rostrumand carina. Seven
individuals are represented by these remains.
Holotype.—The complete specimen (text-fig. 1, C-F), Museum
of Practical Geology, register number 3204.
Horizon and locality.— Lower Senonian, Upper part of J. cor-
testudinarium-zone: Slines Oak Pit, Worms Heath, Woldingham,
Surrey. ?@Same horizon: Watford Tunnel, Hertfordshire.
*° 1889. Mem. Geol. Surv., Geology of London, vol. i.p. 77; 1904. Mem. Geol.
Surv., Cretaceous Rocks, vol. il. p. 232.
nga
+ vineulum, a bond.
A REMARKABLE NEW CIRRIPEDE. 947
Text-figure 1.
A. Verruca prisca J. Bosquet. Upper Senonian to Danian: Europe. Rostro-
carinal view showing the movable opercular valves, which are on the right
side.
B. Opposite view of same showing the fixed scutum and tergum.
C. Proverruca vinculum, gen. et sp.n. Lower Senonian: England. Restoration
of specimen below (fig. D),and from almost the same position.
D. Proverruca vinculum, gen. et sp.n. Lower Senonian, ? zone of Micraster cor-
testudinarium: Watford Tunnel, Hertfordshire. Coll. Mus. Pract. Geol.,
No. 3204. A complete shell in which the movable scutum and tergum are on
the right side and slightly displaced. The rostral- and the carinal-latus are
seen, but the rostral-latus has its upper part broken away showing the lateral
portion of the rostrum below.
E. Opposite view of same showing the fixed scutum and tergum.
F. Same specimen viewed from above to show the broadly oval outline of the shell,
and the semicircular outline formed by the fixed scutum and tergum.
c., carina; ¢.l., carinal-latus ; 7., rostrum ; 7-./., rostral-latus ; s., movable scutum ;
s’., fixed scutum ; ¢., movable tergum ; ¢’., fixed tergum.
All figures * 15 diam.
64*
948 : MR. T. H. WITHERS ON
Measurements.—The complete shell is very minute, its dimen-
sions being :-—
Length (from base of rostrum to apex of carina) 2:7 mm.
“a (from apex 2 - ) 2-1 mm.
Greatest breadth’ .eceeer 2-2-2: ose ee eee ear 1-7 mm.
: height (from apex of tergum to base)... 1-6 mm.
The isolated valves are somewhat larger, and indicate that the
complete specimen is a young individual. Their dimensions
are:
Movable scutum: length (from apex to middle of basal
margin} 2‘8 mm.; breadth (greatest) 1*2 mm.
Fixed scutum: length (from apex to middle of basal margin)
2-7 mm., when complete, ca. 2°9 mm ; length (from apex to rostral
angle) 3:1 mm. ; breadth (greatest) 2°7 mm.
Fixed tergum: length 2-4 mm.; breadth 1:7 mm.
Description of Specimens.
Although the complete specimen is of great importance as
showing the relative position of the valves, little can be seen of
their inner structure. Moreover, the valves are covered exteriorly
by a film of chalk, or may be a secondary deposit of silica, which
serves both to obscure and unduly to emphasize the bead-like
ornament. This is no doubt due to the shell having been pre-
served in a hollow flint, for preservation in flints is apt, especially
in highly ornamented forms, to make the ornament more
pronounced. Owing to its importance it is too dangerous to
attempt to clean this minute and delicate fossil, and it will have
to be studied in its present condition.
The isolated valves vary considerably both in ornament and
structure, but while there is no doubt as to the identity of the
fixed and movable scuta and terga, it is not at all certain that
the rostral- and the carinal-latus belong to this species. In the
complete specimen the rostral-latus is much broken, and the
outer structure of this and the carinal-latus somewhat obscured.
Without being able to disarticulate these valves it is impossible
to compare them with the supposed isolated examples, of which
-even the two rostral latera differ from each other. It is pro-
bable that the variation in ornament is due to varying states of
preservation.
Vhe shell is broadly oval in outline, much elevated, the walls
perpendicular on each side, and the movable scutum and terguim,
when in position, would be inclined at an angle of about 30° with
the base; rostrum and carina at either end, the rostrum wider
than the carina and slightly more developed on the right side, the
space between the rostrum and carina occupied on one side by
the fixed scutum and tergum, and on the other by the carinal-
latus, the rostral-latus for its whole length overlapping the lateral
portion of the rostrum.
Valves highly ornamented, the ornament consisting of strong
A REMARKABLE NEW CIRRIPEDE. 949
transverse rulges, which, where crossed by the longitudinal ridges,
are broken up into bead-like prominences. The fixed and the
movable tergum have this bead-like ornament only on the apico-
basal ridge, the longitudinal ridges being absent on the remainder
of the valve. On the movable scutum the longitudinal ridges are
prominent only near the occludent margin.
Rostrum widely semiconical, wider than the carina, bowed in-
wards, its apex not freely projecting, and the lateral portion
slightly more produced on the rostro-lateral side, on which the
transverse arrangement of the ornament is pronounced.
Carina subtriangular, bowed outwards, moderately convex
ti ‘ansversely, the apex freely projecting ; a short distance from
the carino-lateral margin and extending from the apex, is a
prominent longitudinal ridge formed by the valve being folded
along this line.
Fixed scutwm (Pl. I. figs. 1a, 6) subtriangular, apex acuminate,
considerably convex transversely, with a wide submedian fold or
vidge extending from the apex; on the occludent side of the
ridge the erowth- lines are upturned, and on the tergal side almost
straight ; “oceludent and basal margins convex; tergal margin
concave. On the inner surface a comparatively wide portion of
the valve near the occludent and tergal margins is marked with
growth-lines which meet on a raised ridge below the apex ; this
ridge serves for the reception of the scutal angle of the tergum.
The pit for the adductor muscle takes up a considerable portion
of the lower half of the valve.
Fixed tergum (Pl. I. figs. 4a, 6) subrhomboidal, considerably
convex transversely, with a flat-topped apico-basal ridge which
widens gradually downwards, and on this ridge the transverse
ridges are broken up into small bead-like prominences owing to
the crossing by the longitudinal ridges; basal margin acutely to
broadly rounded. On the inner surface the apical portion is
marked with growth-lines for about one-fourth the extent of
the valve.
Lateral valves—The two lateral valves seen in the complete
specimen from Watford are somewhat broken, and as already
stated, I am uncertain whether the three isolated valves are
identical with them, or whether they belong to the species
described (p. 952) as Scalpellum vimineum, sp.n. The two right
valves, considered to be rostral-latera, are obliquely triangular in
shape and considerably convex transversely. One (Pl. I. figs.
9 a, 6) has the inner surface near the rostral margin marked with
growth-lines to a greater extent than in the other valve (PI. I.
fies. 8a, 6). The latter differs also in the direction of the
transverse ridges on the outer surface, for along a line extending
from the apex near to the rostral margin, the transverse ridges
are not continuous but bend abruptly ‘but slightly upwards and
then downwards to the margin. The right valve, regarded asa
carinal-latus (PI. I. figs. 7 a, b), is subtriangular in shape and very
gently convex transversely. On the inner surface the basal
950 MR. T. H. WITHERS ON
fourth of the valve only was covered by the corium, the upper
three-fourths being marked with growth-lines, which indicate
that the valve freely projected to that extent.
Movable scutwm (PI. I. figs. 2, 3a, 6) an acute-angled isosceles
triangle, slightly bowed towards the tergum, but more so away
from the opposing scutum; basal and tergal margins almost
straight ; occludent margin convex. On the inner surface a
narrow portion of the valve along the occludent edge is thickened,
and a shallow pit for the adductor muscle is situated about the
middle of the valve. On the tergal side the inner edge is raised,
and between it andthe outer edge is formed a narrow groove for
the reception of the tergum.
Movable tergum (PI. 1. figs. 5 a, b, 6) of an elongate diamond
shape, almost flat transversely, with a sharp-edged apico-basal
ridge ; carinal margin formed of two lines making an obtuse
angle; scutal angle slightly protuberant.
Affinities of the Genus.—From a phylogenetic standpoint
Proverruca is by far the most important fossil Cirripede that
has yet been discovered, for it serves in a most remarkable
manner to link up the hitherto distinct families Pollicipedide
and Verrucidee.
The family Verrucide consists of only the single genus Verruca,
but the genus includes some 48 species. Of these, two occur in
the Upper Cretaceous (Upper Senonian and Danian), five are
confined to Tertiary rocks (Miocene and Pliocene), and there are
41 recent species, of which one occurs also in the Pliocene. The
shell of Verruca is very peculiar, since it is quite asymmetrical
owing to the unequal development of the valves. There are six
valves, and Darwin* has shown by tracing the development of
the young shell that they consist on one side of the carina and
rostrum unequally developed on their two sides, on the other of
a tergum and scutum most peculiarly modified and immovably
interlocked to form the ‘ wall” with the rostrum and carina,
and a scutum and tergum in their normal and movable condition
forming the top of the shell. Both Darwin tf and Gruvelt have
shown that at the first period of calcification the valves are
almost symmetrical, but during the subsequent growth of the
shell become more and more unequally developed to form the
asymmetrical shell typical of the Verrucide. It is interesting
also that it appears to be a matter of chance whether it is the
right- or left-hand scutum and tergum that are modified to form
the wall with the rostrum and carina.
Proverruca is of a much more primitive structure than Verruca,
and although the valves are disposed to form an asymmetrical
shell as in Verrwea, with the exception of the inequality in size
of the fixed and movable scuta and terga, the valves have
* 1854. Ray Soc. Monogr. Sub-class Cirripedia, Balanide and Verrucide, p. 498.
1855. Palzont. Soc. Monogr. Foss. Balanide and Verrucidee, p. 41.
T 1854. Ray Soc. Monogr. Sub-class Cirripedia, Balanidew and Verrucide, p. 497.
{ 1905. ‘ Monographie des Cirrhipédes ou Thécostracés,’ p. 170.
A REMARKABLE NEW CIRRIPEDE. 951
undergone very little modification in structure from an ordinary
pedunculate Cirripede. The fixed scutum and tergum have a
greater transverse convexity than in an ordinary pedunculate
Cirripede, but, unlike those of Verruca, they can be neealy
identified. ame fact, they prove the correctness of Darwin’s inter-
pretation of the valves in Verruca as deduced from a study of the
valves of the young shell. As is shown by the right and left
movable scuta and terga, it isa feature of both genera that either
the right or left valves may be developed to torm the “ wall.”
If we take such a pedunculate Cirripede as is included in the
sub-genus Scillelepas of the genus Culantica, we see that the
eapitulum is composed of two whorls of valves, the upper com-
prising paired scuta and terga, anda carina, the lower whorl
consisting of three pairs of latera, a rostrum, and a sub-carina.
There is no upper lateral valve between the scutum and tergum,
although the median lateral valve may be homologous with the
valve that becomes an upper lateral valve in the more specialized
forms of Scalpellum. Now if we imagine the almost equal
development of the rostrum and carina, and the suppression on
one side of the lateral valves, the scutum and tergum would be
allowed to form that side of the wall, and the opposing scutum
and tergum would have to lean over at a greater angle to meet
them. We should then have only to suppress the sub-carina,
the median latus, and the peduncle, to get a form such as
Proverruca. This was evidently the history of the form, and
although Scillelepas may not have been the actual ancestor, it
must have been a form somewhat similar. Proverruca still
retains the primitive structure of the valves, as is shown by the
isolated examples, and the two lateral valves, regarded as homo-
logous with the rostral- and the carinal-latus in the Pollicipedide,
are two that remain of the three lateral valves. It is of much
significance that of these two valves only the carinal-latus really
for ms that part of the wall between the rostrum and carina.
The rostral-latus overlaps for its whole length the lateral portion
of the rostrum, and it certainly seems as though with the
approaching attachment of the lateral portions of the rostrum
and carina, the two lateral valves were on their way to sup-
pression. We have only to imagine their absence, the meeting
of the rostrum and carina, and the development of interlocking
ribs to strengthen the attachment of the valves, te turn Proverruca
into a typical Verruca (text-fig. 1, A, B.)
We see that in the non-attachment of the rostrum and carina,
the presence of two lateral valves, and in the structure of the fixed
scutum and tergum, Proverruca is related to the Pollicipedide,
but more particularly to the genus Calantica Gray. In the
asymmetry of the shell owing to the unequal development of the
valves on both sides, and in one of the scuta and terga forming
the operculum, it is related to Verruca, and it is especially near to
the recent deep-sea species with elevated shells. Proverruca un-
doubtedly represents the ancestral type from which has arisen the
952 MR. T. H. WITHERS ON.
recent group of asymmetrical sessile Cirripedes forming the
family Verrucide, and in its structure clearly shows its origin
from the symmetrical pedunculate forms of the family Polli-
cipedide. 1t presents further evidence that the sessile condition
has been arrived at independently on several different lines of
descent during the evolution of the Cirripedia. The Verrucide
have a phylogenetic history widely different from that of the
Balanidee (sensu lato), and evidence is not wanting to show that
the Balanide also are at least diphyletic. The Chthamaline
have almost certainly arisen from some such form as Brachylepas,
while it is extremely difficult, if not impossible, to derive the
Balanine from that source, or indeed from any form as yet
known.
Family PoLLICIPEDIDS.
SCALPELLUM VIMINEUM *, sp.n. (PI. I. figs. 10-12.)
Diagnosis.—Seutum subtrapezoidal, with no apico-basal ridge,
the apex rounded, the growth-ridges arranged in concentric lines
from the apex; occludent margin forming a right angle with the
basal margin.
Material.—Portions of a right and a left seutum, and a com-
plete right tergum which might or might not belong to the
same species.
Holotype.—The left scutum (figs. 10 a, 6).
Horizon and locality Lower Senonian, wpper part of WM. cor-
testudinarium-zone: Slines Oak Pit, Worms Heath, Wolding-
ham, Surrey.
Measurements.—Length of left scutum, 2°9 mm.; length of
left tergum 1°9 mm., breadth, 1-2 mm.
Scutum.—The left valve (PI. I. figs. 10 a, 6), which is more nearly
complete, has the tergo-lateral portion almost entirely broken
away, and is gently convex transversely. When complete the
valve was subtrapezoidal in outline, and there is no apico-basal
ridge. Apex rounded ; basal margin convex, and forming aright
angle with the lower part of the convex occludent margin. Outer
surface ornamented with a number of concentric ridges termin-
ating each zone of growth, and these ridges are broken up into
bead-like prominences where crossed by the longitudinal ridges.
The middle portion of the valve has the bead-like prominences
much more numerous and crowded. A wide portion of the inner
surface on the occludent side, and so far as preserved on the
tergal side, is marked with growth-lines which meet on a raised
ridge below the apex. A deep pit for the adductor muscle is
situated in the middle of the basal portion of the valve.
Tergum (Pl. I. figs. 124, 6) subrhomboidal, with a delicate
apico-basal furrow ; apex slightly curled towards the scutum,
basal portion narrow and pointed. Upper carinal margin slightly
* vimineus, made of wicker-work.
A REMARKABLE NEW CIRRIPEDE. 953
convex, a little shorter than the lower margin, which is straight ;
occludent margin extremely short, and the scutal angle much
rounded and protuberant ; the valve is depressed near the scutal
angle, and forms a ridge parallel to the margin. Outer surface
ornamented with delicate transverse ridges.
Remarks and Comparison with other Species.—These three valves
were found with those of Proverruca vinculum, already described
(p. 946), but there is no evidence to show that they belong to
one individual. The scutum is ornamented like the valves of
P. vinculum, but it differs so much from the homologous valve of
that species that I refer it to a new species, and to the genus
Scalpellum (sensu lato). The small transverse convexity of the
valve seems to preclude the possibility of its having formed part
of a shell as is the case in Proverruca, and therefore representing
a second species of the genus. Its structure renders it more
probable that it formed part of a capitulum of a pedunculate
Cirripede of the genus Scalpellum, but further information is
needed as to the remaining valves before anything more definite
can be said regarding the relationship of the species.
The scutum differs from that of Proverruca vinculum, mainly
in its much less triangular shape, in the occludent and basal
margins forming an angle of 90° instead of about 45°, and in the
absence of an apico-basal ridge. The tergum differs chiefly in
the presence of a delicate apico-basal furrow, instead of a
prominent ridge.
My thanks are due to Dr. F. A. Bather, Dr. W. T. Calman, and
Mr. C. P. Chatwin for help in connexion with this paper.
EXPLANATION OF THE PLATE.
Proverruca vinculum, gen. et sp. n.
Lower Senonian, Upper part of W. cor-testudinariwm-zone: Slines
Oak Pit, Worms Heath, Woldingham, Surrey.
. Fixed scutum. a, outer view of left valve; 6, inner view.
. Movable scutum. Outer view of a comparatively wide left valve.
. Movable scutum. a, outer view of left valve; 4, inner view.
. Fixed tergum. a, outer view of left valve; 6, inner view.
. Movable tergum. a, outer view of left valve with basal portion broken
away; 6, inner view.
. Movable tergum. Outer view of smaller but more complete left valve.
. Carinal-latus. a, outer view of right valve; 4, inner view.
. Rostral-latus. a, outer view of incomplete left valve ; 6, inner view.
. Rostral-latus. a, outer view of nearly complete left valve; 6, inner view.
Fig..
Co OC NI OD Ore co bo
Sealpellum vimineum, sp. n.
10. Scutum. a, outer view of left valve of which the tergal portion is
broken away ; 6, inner view.
11. Scutum. a, outer view of fragment of a right valve (portion near rostral
angle) with very pronounced ornament; /, inner view.
12. Tergum. a, outer view of complete right valve; 6, inner view.
All figures X circa 8 diameters.
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PZ. 8. 1914. Pottss PE VE
PHYLLOCHAETOPTERUS ANGLICA.
ON POLYCHAETA FROM THE N.E, PACIFIC. 955
51. Polycheeta from the N.E. Pacific : The Cheetopteridee.
With an Account of the Phenomenon of Asexual Repro-
duction in Phullochetopterus and the Description of Two
new Species of Cheetopteridee from the Atlantic. By
F. A. Porrs, M A., Fellow of Trinity Hali, Cambridge,
and Balfour Student of the University *.
[Received May 30, 1914; Read November 24, 1914. ]
(Plates I-—VI.7 and Text-figures 1-13.)
INDEX.
Page
Introduction ......... Raa nsasiaceciae MAL Re GOD
Diagnosis of Family anal Table of Gane Mek nes eee OOO
Mesochetopterus, gen. n.
IDrEeANOSaS Oe Creams Bal INGOTS coooncaanosenoocoscasansueadsonsaugaeod SETl
MENGar LOTtuSPeNee -osene Coenen eee ss oe eo aks ean Nae cst t eden OS
BY Ee DU CORA LD S| Den Oe SRO nae San pod ban ChURE a ee EOrp: Ree ena Eo eee Ce naa ced 963
The Position of Mesochetopterus in the Family .................. 967
Telepsavus G. Costa.
Relepsauussspey ayseey ccc see ence one antics sec dae ee eevaceeniesoy LOOS
Phyllochetopterus Gr.
P. prolifica, sp. n. (with an account of asexual reproduction
in the species) ......... BeoontLE cen
Some Points in the Woupholosy of ihe hetoptaiias ceAedess 981
Description of a new Species (P. anglica) from British Waters,
and a Comparison of those ee. which form True
Colonies ....... 984,
Remarks on the Gener anise fopten US serail ep, lnctates
LCE TULS Mee nee a ae cay ett Mn 7d) 2, Aetna ie thSoklees tesa Meee OOO
INTRODUCTION.
The Cheetopterids found in the coastal waters of the Gulf of
Georgia and Puget Sound, though the species are few in number,
are very widely distributed and remarkably interesting in their
biology and morphology. My observations were made on three
species, all of which appear to be new :—
Mesochetopterus taylori, gen. et sp. n.
Phyllochetopterus prolifica, sp. 0.
Telepsavus sp.
Of these Mesochetopterus partly bridges the gap between
the remarkable form Chetopterus and the other members
of the family. Phyllochetopterus prolifica possesses a type of
asexual reproduction which is now described for the first time
in the Polychetat. T'elepsavus is a genus hitherto only known
* Communicated by the SecRETARY.
+ For explanation of the Plates see p. 993.
{ A preliminary note on this phenomenon was published in Rep. Brit. Assoc,
Ady. Science, 1912, Dundee, p. 513.
956 MR. F. A. POTTS ON
to occur in the Mediterranean and the Red Sea. In previous
collections the Chetopterids appear to have been represented
only by empty tubes, and their interest has remained unsus-
pected (cf. Johnson, “The Polycheta of the Puget Sound Region,”
Proc. Boston Soc. Nat. Hist. vol. xxix. p. 386).
The collections, of which the worms described here form a
part, were made in the summer of 1911, while I was a guest at
the Biological Laboratory at Departure Bay, Vancouver Island,
which is “maintained by the Dominion Government. I should
like to express my heartiest thanks for the hospitality extended
to me there. The tidings of the death of the Rev. G. W. Taylor,
the first Director of the Station, reached me here last year (1912),
and I wish to place on recor d some slight tribute to the memor y
of one of the pioneers of marine biology i in British Columbia. His
enthusiasm for the study of the vich fauna of the Pacific Coast,
and the patient care which he bestowed upon its investigation
are worthy of great praise. I only knew him in the last year of
his hfe, during a time when, crippled by paralysis, he suffered
greatly, but his kindness and thoughtfulness will always remain
a pleasant memory to me,
In connection with my work in Canada, I wish, too, to grate-
fully acknowledge my ‘indebtedness to the Managers of the
Balfour Fund, who made me a special grant to assist in defraying
the expenses of the j journey.
Of the other forms which are described in this paper, J/eso-
chetopterus minuta was found amongst the collection made by
Mr. Cyril Crossland in the Cape Verde Islands during July and
August 1904. I am much obiiged to him for permission to
describe-this form, and for his kindness in reading through this
paper: Lastly, the new species of Phyllochetopter us, which I
have found to be an. inhabitant of British waters, was obtained
while working at the Laboratory of the Marine’ Biological
Association at Plymouth in the spring of 1913.
Family CuatoprerRtp# Audouin and Edwards.
Polycheeta inhabiting a tube of parchment-like consistency and
very closely adapted to their tubicolous life. Body divided into
two, or sometimes three, distinct regions. Prostomiwm small, often
bearing eyes; peristomium forming a collar, with two tentacles
more or less developed. The first (anterior) region is composed of
a small and fairly constant number of segments; of the two
divisions of the parapodium the notopodium only is developed.
The segments behind this region have biramous parapodia ; the
variation of the notopodia here affords the chief method of differ-
entiating between the genera of the family. Generally two distinct
types are successively met with in the same animal, thus enabling
us to distinguish second (median) and third (posterior) regions.
Throughout the body the notopodia carry capillary sete ; in the
fourth segment one or more are much stronger and thicker than
POLYCH ETA FROM THE N.E. PACIFIC. 957
the rest. Hach neuropodium consists of a double ridge, with several
rows of uneini. The dorsal surface is greatly flattened in the
anterior region, and carries a median ciliated groove, which runs
the whole length of the body in some forms, but is interrupted
in others.
Table of Genera.
(a’) All segments behind anterior region similar.
(6’) Notopodium of posterior segments unilobed
(6) Notopodium of posterior segments bilobed
(a’’) Median and posterior regions both present.
(c’) Segments im median region with bilobed foliaceous
notopodia, each carrying several capillary sete.
(d’) Number of segments in median region variable
(d’’) Two seements in median region ......
(c3) Segments. in median region typically with unilobed
‘notopodia. Peristomial collar well dev eloped.
(e’) Median segments two or three in number; noto-
podia all conical in shape; tentacles long ...... Mesochetopterus.
(e’’) Median segments five in number ; first with sepa-
rate aliform notopodia; others with notopodia
fused to form fans or suckers ; tentacles short...
AS See eich Ranzania
Telepsavus.
Phyllochetopterus.
Spiochetopterus.
Chetopterus.
MESOCHAETOPTERUS, gen. n.
Chetopterids with a well-developed peristomial collar and «
pair of long peristomial tentacles. Body divided into three
regions. The anterior contains 9-13 setigerous segments ; th
parapodia are represented by short and conical notopodia with
capillary sete; in the fourth setigerous segment several of the
dorsal setee are enlarged. The median region is composed of
2 or 3 elongated segments, forming dorsally a flat region, with
continuous lateral borders, covered with glandular epitheliwm
and ornamented with transverse ridges. Typically the noto-
podia are rather enlarged, conical, and fleshy, with a groove
running down the inner border; the newropodia are single in
the first, double in the succeeding segment or segments, and con-
tain wuneint. The posterior region contains a large number of
segments similar to those in Cheetopterus, but with much shorter
notopodia. A dorsal ciliated groove runs from the mouth along
the median line to the posterior end. In one or more of the
median segments the lips are enlarged to form a fleshy organ.
The genus thus agrees with Chetopterus in the reduced number
and specialised character of the segments of the median region.
It resembles Phyl/ochetopterus in the continuous ciliated groove
and the long tentacles.
Diagnoses of Species of Mesocheetopterus here described.
M. taylori, sp.n.—A long but slender Mesochetopterus, living in
a long narrow unbranched tube of opaque parchment, embedded for
the most part vertically in sand and ending blindly. Prostomiwm
very small, without eyes, entirely surrounded and hidden by the
well-developed peristomial collar. The anterior region coniains 9 or
958 MR. F. A. POTTS ON
10 setigerous segments. The median region is composed of
3 segments; in all, the notopodia are of the type described in
the diagnosis of the genus. The posterior region contains @
large number (about 60) of segments; the short notopodia each
with several capillary sete.
Locality. Pacific coast of North America.
MM. minuta, sp. n.—A very small slender Mesochetopterus, living
in tubes of a translucent horny material coated with coarse sand.
Prostomium large and conical ; peristomial collar well developed,
but not so complete as in M. taylori. Just external to the tentacles
is a pair of eyes. The anterior region contains 10-13 segments.
The median region is composed of 2 segments; the first pair of
notopodia are small and clavate, the second pair are of the type
described above for the genus. The ciliated groove is expanded
into a cup in the middle of the second median segment. The
posterior region is composed of segments which are double ante-
riorly, single posteriorly ; each notopodium has a single seta.
Locality. Cape Verde Islands, Atlantic; Torres Straits, Pacific.
MESOCH&TOPTERUS TAYLORI, sp.n. (Plates I., ITI., figs. 5, 6, 9;
Text-figs. 1-5.)
Occurrence and Habits —This animal was first found in De-
parture Bay, near Nanaimo, Vancouver Island, on a wide stretch
of sandy beach, which was partly bare and partly covered with
beds of Zostera. Over the whole area, from the middle of the
beach to the lowest tide-mark, there were to be found brown
tubes about a quarter of an inch in diameter, lined with a brown
parchment-like material, while the outer layer is membranous
and coated externally with sand-grains. The tubes, which pro-
ject very slightly above the surface, are not U-shaped as in
Chetopterus, but go straight down through the sand generally
for about eighteen inches. On reaching the shingle underneath
they sometimes turned and ran horizontally. In one case, where
especial care was taken to obtain the tube whole and uninjured,
it was found to end blindly in a neatly rounded apex. The last
art was much thinner, without the parchment lining. The
total length of the tube in this case was three feet. It is, as a
rule, however, very difficult to obtain the entire tube, owing to
the fact that the sand is deeper in most places and the tube runs
vertically through its whole extent. But without obtaining the
whole tube it is almost impossible to examine entire speci-
mens of its inhabitant, which rapidly retreats to the depths of
its dwelling as soon as the spade strikes the sand. Most of the
individuals collected consisted only of the first two regions.
Later the animal was dredged in two or three fathoms of
water at Nanoose Bay, and also observed on sandy beaches,
between tide-marks, at Victoria on the south end of Vancouver
Island and Olga in the San Juan Archipelago, just over the
POLYCHEZTA FROM THE N.E. PACIFIC. 959
International Boundary. It is thus widely distributed in the
neighbourhood of Puget Sound and the Gulf of Georgia.
One whole specimen was obtained with the following dimen-
sions :—
Length 28:5 em, Width in broadest part 1 em.
Anterior region with 9 segments 1°8 em. long.
Median region with 3 segments 4:0 cm.
Posterior region with 68 segments 22°7 cm.
This was probably a small individual, since in others measured
the anterior and median regions exceeded the figures given
above.
Mesochetopterus is a longer and much more slender worm than
Chetopterus variopedatus. The delicacy and transparency of the
integument, which is so marked a feature of the last-named
species, is not characteristic of the new genus. The whole
surface is a creamy white or yellow, relieved by dashes of choco-
late pigment on the peristomium and tentacles alone. In the
posterior region the dark green gut shows through the body-wall.
The ventral musculature is, throughout the body, more developed
than in Chetopterus.
The prostomium (PI. I. fig. 3) is a small rounded prominence
with unpigmented skin. Itis much more distinct than in Cheto-
pterus. There is no trace of eyes. The peristomiwm forms a
prominent buccal funnel which entirely surrounds the prostomium.
It is, however, shallower, with more gently sloping sides than
in Chetopterus. As mentioned above, there is here a noticeable
development of the chocolate pigment, which does not dissolve in
alcohol, and so is retained by the preserved specimens. The
peristomial tentacles originate just outside the peristomial collar.
They are stout grooved structures (very extensile in the living
animal) and measuring, even in their contracted state, 3-4 em.
They contain a pigment similar to that mentioned above. The
mouth is situated between the prostomium and the ventral lip
of the peristomium, and is bordered in some specimens by two
rounded lips below.
The anterior region is convex on the ventral side, concave on
the dorsal. It differs, however, generally from that of Chatopterus,
firstly in the narrower width and secondly in the shortness of the
notopodia. In 21 specimens examined, 10 had 9 segments, an
equal number had 10 segments, and a single individual 11. In
Chetopterus variopedatus, similarly, though 9 is the typical
number, Joyeux Laffuie observed indiv iduals with 10, 11, and
even, in one case, 12 segments. The parapodia of the anterior
region are all similar and represented by the conical notopodia,
which increase slightly in size as we pass posteriorly. The
dorsal curvature is not so distinct as in Chetopterus. ‘The last
segment does not bear an appendage representing the neuro-
podium, such as occurs in C. variopedatus (Joyeux Laftuie, 9,
p. 257, pl. xv. fig. 2) and other species.
960 MR. F. A. POTTS ON
In the notopodium of Chetopterus there are two long straight
rows of capillary sete, embedded for the greater part of their
length in the parapodium, but projecting from the surface for
short but equal distances. They are all lanceolate in type, but
the dorsalmost sete differ from the rest in being slender and
scarcely dilated. This tendency to differential development. is
greatly exaggerated in the 4th segment. The dorsal sete
preserve their slender lanceolate character, but a number of
the ventral sete become short, strong, truncated at their ex-
tremities, and black in colour. Those situated most ventrally
present these characters in the highest degree (text-fig. 4 A).
In Mesochetopierus the sete (Pl. ILI. fig. 9) are more delicate
than those of Chetopterus, aud have a different arrangement, owing
to the shortness of the parapodia. They are mostly of a distinct
lanceolate type and equal in length, but a few of the dorsalmost
are much longer and more slender, projecting in a ragged tuft
from the extremity of the neuropodium. ‘They occur in a slightly
curved row. In the fourth parapodium there is a single line
making a very pronounced curve. Ventrally there are about
12 modified setz.
Median Region.—Though consisting of three segments only, it
is much longer than the anterior region. In Chetopterus the
median region is characterised by the transparent nature of the
body-wall and the reduced width of the segments; here every
segment is of a uniform width greater than that of the anterior
region, the dorsal surface is flattened and glandular, and enclosed
on each side by a continuous border formed by the thin upturned
edges of the segments (PI. I. fig. 2, fr.), and the ventral muscles
are much larger and stouter than in Chetopterus.
When the animal is alive and inside its dwelling the lateral
borders are approximated so as to form an imperfect tube
dorsally. Their appearance in text-fig. 1 does not do justice to
their extent in the living animal.
The parapodia in Mesochetopterus have not undergone the
great and diverse modification occurring in Chetopterus (compare
text-fig. 1, A, B). They resemble very closely those of the abdo-
minal region and attain a similar development in all three
segments. The notopodia are short and conical, little larger
than in the abdomen; they possess an internal skeleton of two
or three stout capillary sete. They are on the distal surface
and approximated to the middle line. On the inner surface
they possess a ciliated groove which meets the median groove.
T think that there is little doubt that they act as accessory organs
for the collection of microscopic food, interrupting the dorsal
channel, and separating food-particles from the current by the
action of the cilia contained in these grooves. The neuropodia
are slightly different in the three segments. The ventral surface
of the anterior region (PI. IIT. fig. 5) is entirely oceupied, as in
Chetopterus, by a “plastron” (pl.) with a slightly wrinkled
appearance to the naked eye and composed of high epithelial
POLYCH ATA FROM THE N.E. PACIFIC, 96]
Text-figure 1.
A. Chetopterus variopedatus. B. Mesochetopterus taylori.
Anterior view of median segments.
al., alimentary canal; cil.gr., dorsal ciliated groove ; fr-, lateral borders ; m2z.,
ventral longitudinal musculature; new., neuropodium ; not., notopodium
with its acicular setee.
cells. From this, along the median region, there runs a wedge-
shaped prolongation (w.), which narrows down in the second
segment to a median groove. But in the first segment it is
broad, and the neuropodia are restricted to two narrow lateral
strips of different appearance. Here, then, each neuropodium
is a single structure ; in the two succeeding segments the neuro-
podia are much broader, extending almost to the middle line,
and are divided into a dorsal and a ventral half, the former being
slightly smaller and pushea a little forward. Hach contains
several rows of uncini. There is no indication whatever of the
fusion of the neuropodia to form a sucker-like median structure,
which occurs in Chetopterus. A typical uucinus of this region is
figured in text-figure 3.
The dorsal surface of the median region is covered by glandular
epithelium, raised into transverse furrows. ;
Proc. Zoou. Soc.—1914, No, LXV. 65
962 MR. F. A. POTTS ON
Posterior Region (text-fig. 2).—In the one complete animal
which I obtained this contained 64 segments, a much larger
number than has ever been observed in Chetopterus (Joyeux
Text-figure 2,
not
A. Chetopterus variopedatus. B. Mesochetopterus taylori.
To show difference in form of typical segments of the posterior region.
cir., cirrus-like appendage of the neuropodium. Other lettering as in text-fig. 1.
Text-figure 3.
Mesechetopterus taylori. X 325.
Uncinus from first segment of median region.
POLYCH&TA FROM THE N.E. PACIFIC. 963
Laffuie, 9, gives 27— -40). The segments differ from those of the
median region chiefly in their shorter length. The whole region
has the appearance of a string of beads, each segment being
rounded and connected with its fellows only by a narrow neck,
through which run the intestine and the nerve-cord (PI. ILI. fig. 6).
The rounded appearance is due to the very much swollen portion
of the segment which contains the generative organs. The two
notopodia have coalesced for a considerable ‘part of their length,
so that the free portions are very short indeed. In them are
contained several (about 8) capillary setze. The neuropodia con-
stitute a continuous narrow ridge half encircling the segment ; it
bears on each side two uncinigerous tori, w hich are, however, by
no means so well developed and independent as 1s the case in
Chetopterus. The dorsal torus is a little smaller and placed
rather more anteriorly.
‘The small cirrus-like appendage (ci7.) found just outside the
dorsal torus in Chetopterus is not present in Mesochwtopterus.
MESOCHZTOPTERUS MINUTA, sp.n. (Plates IT, IIT. figs. 7, 8;
Text-figs. 4, 5.)
Occurrence.—This species was found twice by Mr. Cyril Cross-
land during his visit to the Cape Verde Islands in August and
September, 1904, once at St. Vincent, and again at Porto Praya.
On the first occasion tufts of tubes were collected on the shore
(?amongst sand) at low tide, and on the second tubes which
contained much larger specimens were found projecting from and
embedded in masses of nullipores. They were associated with
Onuphis and a species of Spionid. In November 1913, I found
this species also at Murray Island, Torres Straits, living in sandy
tubes between tide-marks. Except for their generally smaller
size, these Australian individuals conform with the description
which follows in all essential particulars.
The animal is milk-white in colour, with no pigmentation,
except that of the gut shining through posteriorly. None of the
individuals which reached me was quite complete, but those
from Porto Praya were about 2°5 em. in length, while those from
St. Vincent barely exceeded 15cm. In the largest the maximum
breadth was very little more than 1 mm.
The prostomium (Pl. Il. fig. 4) is a very distinct feature, better
developed than in any species of Phyllochetopterus which I have
been able to examine. It is conical in shape. The peristomiwm
forms a deep cup like that in JZ. taylori, but not so complete
owing to the size of the prostomium, and without the considerable
development of pigment which occurs in the other species. At
the base of the prostomium come off the two long peristomial
tentacles. The eyes were only seen in the specimens from Porto
Praya, in which, however, they were very distinct. They are
curiously placed on the peristomium just outside the bases of the
tentacles.
In the anterior region the number of segments is variable. In
6o*
964 MR. F. A. POTTS ON
the two larger individuals from Porto Praya there were 13 or 14
respectively ; in those from St. Vincent 10 segments in most,
9 in one or two. ‘The notopodia show differences from those 1n
the corresponding region of M. taylori, due partly to the great
difference in size of the two animals. In the Cape Verde species
there is naturally a very much smaller number of setee in each
parapodium (about 20 or 30). The setz themselves are easily
distinguishable from those of JZ. taylori (cf. text-fig. 5) by
their shorter heads. As in that species so here, dorsally they
become longer and more slender, with an almost symmetrical
Text-figure 4.
y
a
——————
A
A. Mesochetopterus taylori, X 70. B. Mesochetopterus minuta, X 325.
Enlarged setze of fourth segment.
lanceolate head. The fourth segment (text-fig. 4 B) carries ven-
trally a number of dark brown modified setz varying from 4 to7.
The larger ones have a distinct likeness to those of J. taylori,
though, of course, they are much smaller. There is a tendency
for the oblique edge of the seta to be fimbriated. This is
apparently due, not to a natural serration, but to splitting of
the fibrous chitin of the seta.
The median region consists of only two segments, but it is
nearly twice as long as the anterior region. It is the great
likeness in the configuration of the median region which shows
POLYCHEZTA FROM THE N.E. PACIFIC. 965
quite plainly that these two worms from the Atlantic and the
Pacific must be placed in the same genus. In both the segments
are broad and flattened, with a glandular dorsal surface raised
into transverse ridges and traversed by a continuous ciliated
groove. In both, too, the parapodia are similar, and there is an
enlargement and incipient modification of the ciliated groove
to form a cup-shaped organ. ‘There are, of course, minor differ-
ences, namely, in the development of the notopodia of the first
segment and in the presence of a rounded lateral border to
the region without the thin upturned edges which are character-
istic of W. taylori.
Text-figure 5.
A B
Typical notopodial setze of anterior region.
A. Mesochetopterus taylori. B. Mesochetopterus minuta.
The first segment is much longer. The notopodia are even
less developed than in JZ. taylori—in fact, they are merely clavate
papille like the second pair of per istomial appendages in Phyllo-
chetopterus. In the individuals I examined there were two or
three slender setze embedded almost entirely in the notopodium,
but projecting very slightly from the surface and ending in a
slight lanceolate head.
The neuropodium is a single structure, separated from its
fellow on the other side by a wedge-shaped prolongation of
glandular tissue exactly like that described above for J. taylori.
The uncini contained in the neuropodium are very similar to
966 MR. F. A. POTTS ON
those of the other species, though much smaller. But they’
have six or seven teeth, or very nearly the number which is found
in WM. taylori.
The second segment is about two-thirds the length of the
first. The notopodia of this segment are exactly like those
described as typical for the genus—conical and enlarged, with a
groove running down the internal border towards the median
eroove. It contains a couple ot siender acicular sete, which do
not project terminally from the parapodium as in the first
segment and have no distinguishable head. The neuropodia are
double structures and call for no remark.
About the middle of this segment there is an enlargement of
the lips of the ciliated groove rather like those occurring in the
second and third median segments of I. taylori. Itis exceedingly
interesting to notice, however, that in some individuals the lips
approximate posteriorly, and an almost complete circular eup is
formed like that in the 13th segment of Chetopterus. This is
a variable character in the species, however—in the individual
figured here there is no posterior fusion of the lips. But from
the variations which occur in the genus Mesochaetopterus we can
undoubtedly see how the accessory feeding-organ in Chetopterus
has arisen from the ciliated groove.
The posterior region (Pl. III. figs. 7, 8) is composed of short
rounded segments. As none of the specimens is complete, I am
unable to say how many are found. Anteriorly each segment
is divided into two by a slight constriction. The proximal half
bears the parapodium. As in J/. taylori, the free part of the
notopodia projects very slightly from the surface. As a rule,
they contain a couple of thin acicular sete, sometimes only a
single one. The neuropodia have the usual double uncinigerous
torus, the dorsal part being very small.
The distal half of the segment is the part which in M/. taylori
is diminished to form the neck between successive segments.
The two species here described differ extraordinarily in size,
for while J/. taylori is the largest Chetopterid known, JZ. minuta
is probably the smallest. I shall have occasion to remark upon
the great variation which occurs in the development of the
prostomium in the genus Phyllochetupterus, but in this respect
these two species differ still more widely. But while WZ. minuta
resembles many species of Phyllochetopterus in the character of
the prostomium, the peristomial collar is deeper and _ better
developed than is ever the case in the latter genus, and the
second pair of peristomial appendages—which are so character-
istic of Phyllochetopterus—are, I think, absent here. These two
circumstances are, I believe, connected. Then, again, the number.
of segments comprised in the median-region is quite constant,
but different in the two species—two in J/. minuta, three in
M. taylorii—and this draws another very definite distinction.
There are other differences, which I have mentioned above.
Some of these, ¢. g. number and shape of sete, are partly dependent
POLYCHHTA FROM THE N.E. PACIFIC. 967.
upon the differences in size of the animals. Speaking generally,
however, these two species are far more definitely distinguishable
than any pair of species in Phyllochetopterus or in Cheetopterus.
No less interesting than the diversity in form of the two
species is their curious distribution. While W/. taylori is so far
only known to oceur in the coastal waters of the N.E. Pacific,
M. minuta has already proved to have a much more extended
range. The type-specimens come from the subtropical regions
of the North Atlantic, but I was surprised to find a Jeso-
chetopterus of common occurrence in Torres Straits, which is,
without doubt, identical with JZ. minuta. It will prove, I
venture to predict, a widely spread Indo-Pacific form, and its
absence from previous descriptions only illustrates the difficulty
of obtaining a representative idea of a Polychet fauna from
general collections. Though J. minuta may be found in the
Tndian Ocean, it is less likely to turn up in the Red Sea and the
Mediterranean, where so much attention has been given to the
obseurer forms of Polychet worms, and the distribution will
probably remain discontinuous in type.
The Position of Mesocheetopterus in the Family.
It can hardly be doubted that a close relationship exists
between Chetopterus and Mesochetopterus. The structural
differences between them correspond closely with the different
kinds of tubes which they occupy. Chetopterus possesses a much
wider tube, in which it fits very loosely, and it is for this reason
that the excessive and bizarre modifications of the median region
have been produced. Adhesive organs are needed to maintain
the position of the worm in the tube, and these are formed by
the fusion of the notopodia giving a cup-shaped sucker. Ciliary
action alone would be too feeble to produce an efficient circulatory
current in so wide a space, and hence the fans of the 14th—16th
segments exist. The notopodia of the anterior and posterior
regions are concerned in the movement of the animal up and
down in its tube. They must be long enough to touch the walls
and so attain to much greater dimensions than in Mesocheto-
pterus. The greater or lesser length of the parapodia causes, as
we have seen above, some difference in the arrangement of the
setie.
Chetopterus possesses, too, a complicated method of feeding,
which is responsible for further ditferentiation in its external
structure. This method has been lately described by Enders in
detail (4). The long aliform notopodia of the 12th segment and
the dorsal cup of the 13th segment alike aid in separating food
from the respiratory current, and compacting it into masses
which are swept forward in the ciliary groove to the mouth.
The middle region, then, fulfils a double function in promoting
the circulation of water in the tube and collecting food, different
segments bemg specialised for each task, In consequence of this
968 MR. F. A. POTTS ON
division of labour, a number of segments is included in the region,
larger than in Jesochcetopterus.
T have not found it possible, in the absence of experimental
observations, to reach any such clear conclusions as to the
functions of the different organs of Mesochetopterus. Food is
partly collected by two long grooved tentacles, which are constantly
projecting from the mouth of the tube and sweeping over the
surface of the sand in search of small fry. Enders makes a
similar observation on another Cheetopterid with long tentacles,
to which he applies the name Spiochetopterus oculatus? He
describes it as scraping with its tentacles the sides of the aquarium
in which it was kept; the diatoms thus dislodged were swept
up the ciliary grooves of the tentacles into the mouth. Nutritive
particles are, no doubt, also collected by the action of the cilia of
the buccal funnel as in Chetopterus, and, lastly, I think, the
notopodia of the median region have a similar function to those
of the 12th segment in the above-mentioned worm. A branch
of the median ciliated groove runs along the inner surface of
each of them, and in my view such minute organisms as are not
strained from the circulating sea-water anteriorly are here
arrested by the parapodia, mixed with mucus secreted by the
glandular epithelium of the surface and swept into the median
groove and along to the mouth.
The ciliated groove in Mesochetopterus, as in Phyllechcetopterus,
is quite continuous in its course from the head to the tail. The
primary function of such a groove is to maintain a respiratory
current through the tube, as is well seen in forms with a trans-
parent dwelling like Phyllochetopterus prolifica.
In Chetopterus, owing to the development of the respiratory
fans, the ciliated groove does not extend farther back than the
13th segment and has changed its function, being now employed
in the collection of food. We may well suppose that in Meso-
cheetopterus the action of the cilia causing a flow of water from
head to tail is not continuous, but is reversed on occasion to carry
food back to the mouth.
Cheetopterus is without doubt the form most specialised in
structure in the family, and this is shown not only by the modi-
fications of the median region but also in the discontinuous
ciliated groove and the shortness of the tentacles. It is impossible
to agree with Enders, who supposes that the long tentacles of
Spiochetopterus “have undergone a considerable specialisation ”
and speaks of the shorter tentacles of Chetopterus as more
primitive than those of Spiochetopterus.
The group of Cheetopterids, which includes Ranzania, Phyllo-
chetopterus, Telepsavus, and Spiochetopterus, ave distinguished by
possessing long tentacles (the most important organs in procuring
food) and a complete ciliated groove. A differentiation of the
median from the posterior region is sometimes not found at all
(Telepsavus, Ranzania). If it occurs, the number of segments
in the median region is very variable (in species and even In
POLYCH HLA FROM THE N.E. PACIFIC 969
individuals), and they differ very little from the abdominal
segments. But the other two points are, without doubt, primitive
characters, and the long tentacles (but not the ciliated groove)
are shared with the Spionids.
Mesochetopterus forms the connecting-link between these
primitive forms and the specialised Chetopterus.
In the possession of long tentacles and a continuous ciliated
groove it resembles Phyllochetopterus. In the development of a
very distinct median region composed of a small number of
segments with grooved notopodia, which possibly assist in the
collection of food, it comes near to Chetopterus.
TaLEPsAvus Gabr. Costa.
Only two species of this genus have been described hitherto:
these are 7’. costarum Claparéde, from Naples (1), and 7’, bonhouret
Gravier, from Djibouti in the Red Sea (5).
TrLEpsavus sp. (Text-figs. 6-8.)
Though this Polycheet is a very common and widely distributed
member of the beach fauna of British Columbia, my collection
contains only one specimen which is at all well preserved, and in
this the head is incomplete. Such individuals as were preserved
in their tubes were quite worthless, owing to the impermeability
of the material, which thus differs from that formed by Phyzllo-
chetopterus considerably more than their appearance seems to
show. The following account of the species is thus very incom-
plete, and I refrain from giving a name until the species can be
better defined. I should also like to state my opinion that the
validity of Gravier’s species is questionable until an actual
comparison of the Mediterranean and Red Sea forms is made.
Occurrence.—On the same sandy beach at Departure Bay,
which I have described in my account of MJesochetopterus, the
tubes of a second smaller Cheetopterid were discovered. They
are composed of a translucent horny material and are annulated,
the joints occurring at short intervals. In length they some-
times exceed a foot and a half, running vertically down through
the sand and ending in a neatly rounded apex. In diameter the
tube is less than 2 millimetres, the worm fitting fairly tightly
within its habitation. A single individual occurs in each tube,
with its long peristomial tentacles often projecting from the
aperture.
The distribution of Zelepsavus has been found to be a wide
one. Generally it may be said to occur wherever Mesochetopterus
exists. I have collected both together in Departure Bay, at Olga
in the San Juan Archipelago, on Ballard Beach, Seattle, U.S.A.,
and by dredging in two or three fathoms of water at Nanoose
Bay, Vancouver Island. :
At the end of August 1911, too, I found Zelepsavus at Skide-
gate in the Queen Charlotte Islands, 500 miles to the north,
living in muddy gravel. Here and at Departure Bay at the
970 MR, F, A. POTS ON.
beginning of the month the female worms had well-developed
orange gonads.
Size.—The almost complete individual measured was about
5 em. long and nearly 2 mm. broad (i. e. about the same length
as and rather broader than the two species hitherto described).
T cannot, unfortunately, say anything about the structure of
the head. The anterior region consists of 9 segments. In my
specimen the 7th, 8th, and 9th segments, but particularly the
two latter, are much longer than the others. In this particular
the Canadian form agrees with the description of 7. bonhowuret,
Text-figure 6.
Ms
|
B (ae
Telepsavus sp.
A. Enlarged seta from 4th segment. X 70.
B, C Notopodial setie from 5th segment. A lanceolate type occurring in
H a dorsal position. X 325.
while the figure of 7’. costarwm shows little of such a differ-
entiation. I should like, however, to assure myself of the
importance of this point (which is strongly emphasised by Gravier)
on more and better material and by a comparison with specimens
from Naples.
In the 4th segment there is a single greatly enlarged seta
(text-fig. 6 A), The point of this is an irregular triangle. The
base is slightly curved and quite entire (in 7. bonhourei it is
furnished with a fringe of projecting points). The longer of the
outer sides is furnished with very unequal serrations. There is
also an auxiliary bundle consisting of a few fine sete, such as
is figured for 7. costarum, while it is expressly stated that it
does not exist in 7’. bonhowrei. A series of types of notopodial
setxe 1s Shown in text-fig. 7.
971
POLYCH ZETA FROM THE N.E. PACIFIC.
Text-figure iG
\
Telepsavus sp.
Other types of notopodial sete from anterior region. X 820.
Text-figure 8.
mot, acc.
“pap.
Telepsavus sp.
Lateral view of posterior region.
obe of this; pap., papilla of unknown
nature; new7., neuropodiun.
not., notopodium ; aec.l., accessory ]
Behind the anterior region there are between 30 and 40 seg-
ments, all of the same type with a pifid notopodium (text-fig. 8).
The first three segments are distinctly longer than the rest, and
972 MR. F. A. POTTS ON
there is no such development of glandular papille on their dorsal
surface as is figured in the Neapolitan species (but absent in
1’. bonhouwrei). ‘he parapodia throughout consist, as is usual, of
a bifid notopodium with capillary sete, a lateral accessory lobe
such as occurs in Phyllochetopterus, and a double neuropodium
consisting of two adjacent uncinigerous ridges. There is, more-
over, in all the segments a very distinct, occasionally double,
papilla just anterior to the notopodium. ‘This may be the
nephridial papilla, but, if so, its position is more dorsal than
usual.
The ventral surface of the anterior region is deeply tinged
with a brown to purple pigment. In thisit apparently resembles
T. costarum, while in 7’. bonhowrei the 7th segment alone is
markedly pigmented.
From the points which have been stated here it will be seen
that the form certainly differs specifically from 7’. bonhowrei and
probably also from 7’. costarwne.
PHYLLOCH EroPrEeRUS Grube.
PHYLLOCHATOPTERUS PROLIFICA, Sp. n. (Plates 1V., V.; Text-
fig. 11.)
Phyllochetopterus of small size (1-3 em. in length), with eye-
spots. Anterior region usually with 12 setigerous seyments; a single
enlarged seta in each parapodium of the 4th segment. Median
region with a very variable number of segments. In segments of
posterior region each notopodium contains a single seta. Tubes
creeping, usually containing several individuals and possessing
several short branches opening to the exterior.
This species of Phyllochetopterus was first collected outside
the harbour of Nanaimo. Here, as was proved by frequent
dredgings between the Five Finger Rocks on the north and the
island of Gabriola to the south, the muddy bottom is covered
with a thick growth of hexactinellid sponges (Bathydorus dawsoni,
Aphrocallistes whiteavesii), and associated with these are the thin
and delicate tubes which prove to contain Phyllochetopterus,
sometimes sparsely scattered, sometimes in such thick and
tangled masses that the dredge contained little else. Later in
the year, a visit to the Marine Biological Station of the Uni-
versity of Washington at Friday Harbour showed that this
annelid oceurs abundantly in various localities in Puget Sound.
It was dredged at many points in the San Juan Archipelago
and found associated with very different companions. In the
dredgings from deeper water (down to 60 fathoms) masses of
large barnacles (Balanus aquila) and the tubes of the Polychet
Sabellaria also occurred; in shallower waters (up to 5 fathoms)
the Phyllochetopterus tubes were entwined with red seaweeds.
But though the vertical range as seen above is fairly notable, it
never occurs above low-tide mark, where it is succeeded by the
other members of the family, MJesochetopterus and Telepsavus.
POLYCHETA FROM THE N.E. PACIFIC. 973
The fact that though so abundant this Phyllochetopterid has
remained up to the present undescribed is due to the slender
nature of the tubes, which easily escape identification as the
habitation of an annelid, and to the difficulty of preservation.
In its wide distribution and its habit of forming intertwined ~
masses of tubes, P. prolifica resembles P. socialis of the Mediter-
ranean, of which Claparede wrote in 1866: “Cette annélide est
probablement l’espece la plus abondante dans le golfe de Naples*,
ou ses tubes juxtaposés, grisatres, papyracés et enchevétrés les
uns dans les autres par leur extrémité postérieure, paraissent
former d’immenses prairies. Du moins les pécheurs apportent-
ils & premicre requisition, sous la nom de ceppa grande, des piéces
quon prendrait pour de grands quartiers de gazon, et qui sont
formées presque exclusivement par les tubes de ce Phyllo-
chétoptere.” In P. pictus, too, described by Cyril Crossland (2)
from Zanzibar in 1903, a similar habit was observed. On the
occasion of its discovery the tubes ‘‘ were found clustered together
in considerable numbers on the underside of a large stone at
low-water level.” But in the remaining species of the genus
hitherto described the worms inhabit straight solitary tubes.
As will be seen later, examination of the individuals of
P. prolifica shows that the size and external characteristics of the
species tally fairly well with the description of P. socialis. But
that there is a deeper physiological connection is indicated by the
occurrence in both species of more than one individual in a single
tube. The phenomenon of asexual reproduction, which Claparéde
suggested as the cause of the colonial habit in P. socialis, 1 wish
to record here as the outstanding feature of P. prolijfica.
I will first give Claparéde’s statement on this point which
follows immediately after the passage quoted above :—‘* L’étude
de cette annélide a fait surgi quelques curieux problémes physio-
logiques. Les ceppe grandi qu'apportent les pecheurs sont formées
exclusivement par des individus d’un méme sexe, généralement
des males, les femelles étant a ce qu’il semble beaucoup plus rares
que les males. Hn outire, chaque tube est réguliérement habiteé
par deux ou trois individus, tous adultes et mtrs. Le tube est
cependant si étroit que seul lindividu antérieur peut fait sortir
ses tentacules par l’ouverture, tandis que les suivants sont empri-
sonnés derriére lui. Dans de paveilles conditions, on doit sup-
poser tout naturellement que ces derniers ont été engendrés par
bourgeonnement postérieur a lextrémité du premier, et que peut-
étre méme tous les individus d’une méme ceppa sont nés par
gemmation. ‘Toute-fois je n'ai pas réussi a vérifier l’exactitude
de cette hypothése. Je ne suis pas méme trés-certain des rapports
des tubes entre eux. Ces petites habitations cylindriques larges
A peine d’un millimetre et longues parfois de 8 a 10 centimetres,
sont irréguliérement contournées dans leur partie postérieure,
sondées les unes aux autres, et ne peuvent se séparer sans
* T understand that P. socialis is now considered something of a rarity at
Naples.
974 MR. F. A. POTTS ON
déchirures. I] m’a semblé quelles s’anastomosaient parfois, ce-
pendant j’éprouve quelque hésitation & affiumer ce point. Ilya
done, on le voit, encore bien des questions 4 vider au sujet de ces
vers.”
The incompleteness of Claparéde’s observations leaves us in
some uncertainty with regard to P. socialis. As I understand
the foregoing passage, he was unable to assure himself as to which
.of the following alternatives was correct. Either ;
(1) There are a number of unbranched tubes packed close
together, but without connection, each tube containing two or
three individuals, but having only a single opening ;
Or else (2) these adjacent tubes are in reality connected with
one another, and the whole bundle of tuhes (ceppa) is a single
colony, all the worms contained in which have probably been
budded off from a single original individual. This is supported
by the fact that sne worms in a bundle are all of the same sex.
In P. prolifica, on the other hand, the relations of the tubes
are perfectly clear. Where they come into contact they do not
adhere in the complex manner described by Claparéde. In the
following paragraph I state shortly the conditions which are
found in the colonies.
The tubes are comparatively long and sometimes divide into
two or more branches of approximately equal length. Most
contain more than one worm and some as many as six. The
main tube is provided with several short branches which open to
the exterior, but the number of openings does not correspond to
the number of worms in the tube. The worms can change their
position in the tube fairly rapidly and can turn round and pass
each other. Those individuals which occupy a favourable position
protrude their long tentacles from one of the openings to assist
in the collection of food.
It occurred to me on first observing the above facts, while I
was still unaware of Claparéde’s observations on L. socialis, that
an explanation was probably to be found by supposing some sort
of asexual reproduction to occur in the species. The nature of
this reproduction was indicated by a discovery made during an
early examination of the living material at Departure Bay.
T was surprised to find emerging from freshly dredged tubes
very short individuals which differed considerably from the
normal forms in the constitution of their bodies. One of these
only measured 6 mm., which is +—7 of the normal length. The
3 =
segments of which it was composed were arranged as follows :—
Anterior region ...... 8 segments.
Median LS Vy Fa Bk 2 es
Posterior 2 hee 20-30 ,,
Not only did the anterior region consist of less than the
normal number of segments (12), but also there was no segment
with a modified seta such as occurs in all adults of the species,
and the peristomial tentacles were represented by minute stumps.
The number of segments in the median region was exceedingly
POLYCH STA FROM THE N.E. PACIFIC. 975
small, but the segments themselves both here and in the posterior
region were normally developed, while there could not be the
slightest doubt but that the anterior region was in process of
regeneration.
A second individual found at the same time gave another stage
in the phenomenon. It was more nearly the normal size. The
regions contained the following numbers of segments :—
Anterior region...... 11 segments.
Micdiangy sy) sack : 5 P
Rosteniony sone... 20-30 __,,
The anterior region thus contained nearly the normal number
of segments, but it was shorter than usual and the peristomial
tentacles were less than half the usual length. ‘This seemed a
second case in which the anterior region was repeneratine.
On returning to Cambridge, a detailed examination of a large
amount of material preserved in formalin was made. Tube by
tube was taken and slit up, and the length and constitution of
the three regions noted in all the individuals contained therein.
Jt was quite clearly shown that living constantly though the
animals do within the shelter of a tube, regenerating specimens
are frequently found. From an examination of these it seems
certain that a fragment of Phyllochetopterus containing only
segments of the median and posterior regions easily regenerates
an entire anterior region. The number of cases in which this
phenomenon has apparently occurred, and the fact that in their
protected situation regeneration after external injury is an
unlikely event, leads me to suppose that these animals have the
power of autotomy and that it is exercised for the distinct
purpose of reproduction.
The tables which follow present the results of the detailed
examination of several tubes, each represented by a separate
TABLE L.
Constitution of the three body regions.
A. B. C. | Total length
Anterior. | Median. Posterior. | of worm.
| |
Nas ie 2a 12 | 6 5 | 13mm.
Piece LS ak be No ee a es
INO; Brae cor 12 a none | 18mm,
a | |
(| Small regenerating stump | ?) |
' 1 mm. long; segments | 5 39 |
va e og |
|
|
No. 3 a || not sufficiently differ- \
entiated to count.
976 MR. F. A. POTTS ON
table. The figures in the columns A, B, and C indicate through-
out the numbers of segments in the regions referred to.
In this tube (Table I.) from Departure Bay Nos. 2 and 3 appear
to be complementary, and represent an individual broken in two
in the median region. While, however, the median segments of
No. 3 have already regenerated a short and undifferentiated
region, those of No. 2 have not yet attempted to form a posterior
region. The regenerating end of No. 3 is figured in Pl. V.
fig. 12, and it will be seen that a number of segments have
been marked off by furrows, but that parapodia and sete have
not yet developed. The peristomium is indicated by the rudi-
mentary tentacles. It will be noticed that almost the full number
of segments is established at once, and apparently development
proceeds simultaneously in each of them.
TABLE II.
|
A. Length of A*. Bb. C. Total length.
|
|
|
| ——
| |
LeNow ics: 11 5°5 mm. (4) 10 12 | 20mm.
pate a ee) ees
| No. 2)... 9 4. nm. (+) 8 22 | 15mm.
INOHoIEE: 12 2mm. (3) 8 40 175 mm.
| 28
| No. 4...| 12 | 4mm. (4) 4 (+number of very 19mm. |
| small segments) |
| sos ake
| No. 5 ...| 12 | 65 mm. (3) 14 | 20 21mm. |
Non Ghee: 12 we 3 20 16 mm.
| t
Another tube from Friday Harbour (Table II.) contained six
individuals, all of which with one exception were well-developed
and complete specimens. It will be seen that while the number
of segments is more constant in the anterior region than in the
other two, its length varies very considerably and bears no
definite relation to the total length of the body, In No. 3
(Plate V. fig. 13) it is extremely short, although it possesses the
usual number of segments, which are of normal width and have
the full development of setze. There is, however, no strengthened
seta in the fourth segment, and the peristomial tentacles are
about half-grown.
In the other individuals the length of this region varies from
4-6°5 mm.
*
he fraction in this column represents the proportion borne by the length
of A to the total length.
fo)
a |
=~]
POLYCH-ETA FROM THE N.E. PACIFIC.
Tasie [I1.
A. Length of A. B. | C. Total length.
ieNiosnlacee 12 6 mm. (+) 8 | 26 , 30mm.
| |
| { i Peas
)No. 2...) 0s as | “6 13 6 mm.
fi ea | veer) oo) pri go thee
a very small regenerating |
+ | 5 99, | 9
| No.3...) stump. | 2 | 12mm.
if
= | —— Seen ae
| | |
| No. 4...| 12 | 55 mm. (4) | 10 | 10 | 24 mm.
| } |
| aca arose oa ee a) ey
\ | | 22
INos See 12 | 6mm. (3) a \+15 very small 27 mm.
segments.
In this colony (also from Friday Harbour) there are three
worms complete anteriorly and two fragments, one of which 1s
commencing regeneration, Of the first-mentioned, however,
No. 4 has probably lately suffered the loss of posterior segments,
as shown by the small number of those remaining and the
moderate total length. No. 5 is interesting from the possession
of a tail of very small posterior segments following others of
normal size, and these must represent regeneration after
autotomy.
From the comparative rarity of regenerated tails in autotomised
worms, it is evident that the anterior region is re-formed much
more quickly than the posterior. With regard to the median
region, there are two facts which seem to show that regeneration
of new segments of this type takes place but rarely. These are:
(1) the great variation in the number of segments (from
4 to 14), even in individuals with well-developed anterior and
posterior regions, and
(2) the absence of segments of two different sizes in the
region, | have, however, examined one animal alive in its tube
in which the normal segments of the median region were
preceded by a single newly formed segment of similar type, but
only about half the size of the others, I cannot say whether the
anterior region was fully formed or not, Cases of regeneration
of median segments then do exist, if but rarely.
The facts incline me to suggest that asexual generation only
occurs successfully when fragmentation takes place in the median
region, that regeneration proceeds both on the anterior and
posterior surfaces of the plane of rupture, however small a
number of segments are left on one side, but that these median
segments usually regenerate anterior or posterior segments, and
Proc. Zoot. Soc.— 1914, No. LX VI. 66
978 MR. F. A. POTTS ON
only segments like themselves when the other two regions have
been completed.
Fragments consisting of segments of the posterior region
alone are, however, occasionally met with, and in one case
(Table Tl. No. 3) a minute regenerating stump was found. At
the same time there is no evidence that complete regeneration
occurs from abdominal segments alone.
Another case may be mentioned in which autotomy had
occurred in the middle of the anterior region and a number of
segments of smaller size were superimposed on the older
segments.
Plate V. illustrates two well-marked regeneration stages of the
anterior region from individuals mentioned above, and they
should be compared with the individual of normal development
shown on the same plate (fig. 14).
REGENERATION IN Cheetopterus.
Since the above was written, | have read the observations of
Gravier on autotomy and regeneration in Chetopterus variopedatus.
When an individual is seized by the anterior part of the body, or
when it is strongly irritated, rupture takes place between the
first and second segments of the median region—this being the
place of least resistance. The anterior fragment can reproduce
all the rest: the posterior has not always been regarded a
capable of regeneration. But Gravier (6) describes and | figures a
posterior fragment, collected at Saint Vaast-la-Hogue, which
Carries a regenerated anterior region ay differentiated but quite
minute. Gravier’s drawi ing (l.c. fig. 2, p. 147) resembles almost
exactly those I have given for P. prolifica. The whole anterior
region hardly equals in length a single original segment, but it
possesses a buccal funnel and per istomial tentacles, and the lateral
border is marked out into segments, 12 conical seta-bearing
notopodia being present on each side. Only in the 4th segment
the special sete are not indicated, and the peristomial tentacles
are unequally developed. There are some irregularities, also, in
the notopodia, but, generally speaking, they are of nearly equal
development. Only the last segment is markedly smaller than
the rest, which thus appear to be marked off nearly simultaneously.
The total number of segments is 12, while the normal number is
only 9. Individuals with 12 segments are very exceptional, and
it is curious to find that in regeneration the maximum number
of segments should be formed.
The first segment of the median region is present, but in a
very rudimentary condition. Noto- and neuropodia are visible,
but it has obviously been formed after the segments of the
anterior region.
If we summarise the phenomenon so far as it is known in
Chetopterus, it may be said (1) that autotomy occurs sometimes
POLYCHEHTA FROM THE N.E. PACIFIC. 979
as the result of an unexpected stimulus*; (2) that, following
autotomy, regeneration will take place from the posterior fragment,
the median region thus giving rise to the whole of the anterior
region, and after that replacing the missing median segment.
There is thus a clear resemblance to the manner of regeneration
in Phyllochetopterus and also a clear minor distinction, the
presence of a definite breaking-point in Chetopterus and its
absence in Phyllochetopterus. Autotomy and a complete type of
regeneration are thus to some extent characteristic of the family
Cheetopteride. In Phyllochetopterus prolifica, however, the occa-
sions on which autotomy takes place are so frequent and regular as
to subserve a definite method of asexual reproduction. But the
nature of the stimuli which cause autotomy, and the question
whether the phenomenon is in any sense under the control of the
animal itself, can hardly be approached as yet.
Some advantage may, I think, be gained by comparing the
cases of regeneration studied in other Polycheta with a differentia-
tion of regions. Ivanow (8) and other authors have made a very
thorough examination of these phenomena in the case of the
Sabellid Spirographis spallanzani. Here there are three regions :
the anterior thoracic with the prostomium, bearing the enormously
developed tentacles, and the first three setigerous segments; the
posterior thoracic, consisting of eight or nine following segments ;
and the abdominal, with an indefinite number of segments. Only
such fragments regenerate as consist of abdominal segments or of
abdominal and thoracic segments. ‘Those containing thoracic
segments only always disintegrate. In regenerating fragments,
the hinder end always produces abdominal segments, and the
anterior end regenerates the prostomium and the three anterior
thoracic segments. The posterior thorax develops later by the
metamorphosis of the most anterior abdominal segments, a
striking change taking place in the characters of the parapodia.
The dorsal uncini are replaced by capillary sete, while in the
neuropodium the capillary setz are replaced by uncini.
A similar phenomenon has been described by Watson (12) in
Potamilla reniformis, another Sabellid. Here, in the regeneration
of the anterior region from abdominal fragments, the prostomium
and one new setigerous thoracic segment only are formed as a new
growth: all the rest of the thoracic segments are formed from
abdominal segments in which a modification of the parapodia
like that described above occurs. It is curious that two regions,
differing from each other so little in morphological characters as
do the anterior and posterior thoracic regions of Spirographis,
should have such a dissimilar method of re-formation.
Though my observations on regeneration in Phyllochetopterus
* In 1913, at Plymouth, I noticed that of a tubful of Chetopterus which had
been brought i in, after being kept on board a trawler for 20 hours or so, nearly all had
autotomised, as a result of “the unhealthy conditions, rupture taking’ place between
the first and second segments of the median region.
66%
980 MR. F. A. POTTS ON
are incomplete, it seems probable that the phenomenon here runs
a very different course. The three regions of the Chetopterid
have no probable connection with those of the Sabellid, nor is
there any reason that they should behave in a physiologically
similar manner. In Phyllochetopterus it seems fairly certain, as
I have pointed out above, that regeneration takes place most
often from fragments containing median as well as posterior
(abdominal) segments, and in all cases the full number of
segments in the anterior region is budded off. But, while in
both Cheetopterids and Sabellids the anterior region is restored
by regeneration, in the former family the median region is not
completely re-formed as is the case in the latter. The number of
segments in the median region is so variable in number that for
this and other reasons I am inclined to suppose that regeneration
does not take place here, or only very slowly. ‘There is certamly
not the slightest evidence, in any regenerating individuals which
I have examined, of a transformation of the posterior (abdominal)
segments into median segments, such as characterises the
Sabellids.
Tuer Possrpitiry oF ASEXUAL GENERATION IN P. pictus.
The case of Phyllochetopterus pictus described by Crossland has
already been mentioned. It will be interesting here to quote
some figures which Crossland gives to show the variability in the
numbers of segments in the anterior (A) and median (B) regions.
Hight individuals were examined. They are indicated by the
Roman numerals running across the page :—
I. II, ite Nye Vie Vie VU) Sve
AP gia 16 as 13 15 12 15 13
Bee’ 9 D 7 5 o 8
Here, too, it will be seen that the median region is much more
variable than the anterior, the numbers ranging from 3—9 for a
very small sample of individuals. In view of this and the fact
that the annelid inhabits clusters of tubes, the relations of which
are difficult to make out, I think it is possible that asexual
generation occurs here too.
MISCELLANEOUS OBSERVATIONS ON THE Habits oF
Phyllocheiopterus prolifica.
Owing to the transparency of the tube, in the younger colonies
at least, it is possible to see something of the movements of the
worm within. It shifts its position in the tube by the alternate
relaxation and contraction of the body; when contraction occurs,
the sete of that part of the body affected are braced against the
sides of the tube. The sete of the abdomen can be definitely
POLYCH.ETA FROM THE N.E. PACIFIC, 981
used as organs of progression, pushing the animal along. The
worms can thus move with some rapidity, and they can also turn
within the tube and even pass each other. When at rest they
generally lie with the long tentacles projecting from an aperture
evidently questing for food. All the inhabitants of the tube may
not be able to obtain such advantageous situations, but from
their activity within the tube it is evident that a fr equent inter-
change of position does take place. The worms do not all lie the
same way, so it is difficult to see how a constant and sufficient
circulation of water can be maintained through the whole tube.
Probably at times the animal is entirely quiescent and the
respiratory current interrupted. In all the animals I observed,
the action of the cilia in the dorsal groove and of the notopodia
of the median region is responsible for a current running poste-
riorly, which supplies not only oxygen but also food, as in the
other Chetopterids. This action is sometimes vigorously supple-
mented by the undulatory movements of the abdomen. This
energetic action—a phenomenon often observed, too, in Cheto-
pterus—is possibly necessary for removing objectionable particles
from the neighbourhood of the body. hn P. anglica and to a
less degree in P. prolifica, 1t may be seen, however, that the
circulation of water is not always thorough, for large sections of
the tube behind worms are blocked by fecal masses, and this
may eventually necessitate the abandonment of the old parts and
extension of the colony.
I have not observed any individuals bearing genital products,
but this is probably due to insufficient examinations and I should
not like to suggest that asexual generation has supplanted the
sexual method. Isolated individuals (like those of Chetopterus)
are not able to manufacture fresh tubes in spite of copious secretion
of mucus, and new colonies must be formed in the first place by
a single individual developed from a fertilised egg, though this,
by fragmentation, gives rise to all the inhabitants of the Rcolony
Claparéde made the interesting observation that, in P. socialis, all
the worms in a bundle of tubes developed genital products of the
same sex, indicating that they were all derived from a single
sexually produced embryo.
Some Ports IN THE MoRPHOLOGY OF THE CH&TOPTERIDE.
Variation in Form of the Notopodia.— Behind the anterior
region of the body both notopodium and neuropodium are
present in the parapodium. In nearly all cases the neuropodium
is stable and retains its character as a double uncinigerous ridge.
In Mesochetopterus, where the first neuropodium of the median
region is single, there is a slight modification, and in Cheetopterus
the neuropodia of opposite sides in the median region are fused
to form a sucker.
The notopodium, on the other hand, is very variable, and it
982 MR. F. A. POTTS ON
may be of some service to arrange the different types in a
tabular form :—
A single lobe without sete ....,.......---..-.-- +22. Ranzania, 16th and all succeeding
segments.
(A single seta ........... Most species of Phyllochetopterus
A single conical lobe } in the posterior region.
with setz.
ie sete ............. Chetopterus and Phyllocheto-
pterus aciculigerus, claparedet
in the posterior region.
Ranzania, 13th and 14th segments.
; (Phyllochetopterus, (A conical or aliform erooved struc:
A bilobed foliaceous | in the median region. | ture (with several capillary
structure with seve-~ Telepsavus, 4 Prete ie bt
ral capillary sete. | all segments behind | C’@topterus, Ist segment 0
median region.
| Mesochetopterus, median region.
The two notopodia of the seg-
ment fused toe form a fan-like
structure.
Chetopterus, segments 3-5 of me-
dian region.
(the anterior region.
The structural types can thus be arranged in a neat series, and
there is some evidence to show the direction in which evolution
has taken place. Both the conical and the foliaceous types are
adapted, the one for progression in the tube, the other for the
respiratory function. But since the median region has been
clearly developed from the posterior, and the conical type of
posterior notopodium is far more widespread than the foliaceous
type (Telepsavus alone), it seems reasonable to regard the conical
notopodium as the move primitive. In this case, Telepsavus is x
specialised form, in which all the posterior segments have de-
veloped foliaceous notopodia and the original type has been lost.
But Ranzania, though the parapodia are unilobed throughout the
body, is not to be regarded as the primitive form from which
the Cheetopterids diverged. The absence of sete from most of the
posterior notopodia and of long tentacles may surely be regarded
as secondary. The 13th and 14th segments are so distinct
from the rest as to almost merit inclusion in a separate median
region.
The Head in the Chetopteride.
In the structure of the head there is also a certain amount of
variation in the family. The prostominm is in most cases provided
with eyes, but it is small in all forms and almost surrounded
by the peristomiwm, which forms a collar. In Chetopterus and
Mesocheetopterus this collar is so complete as almost to hide the
prostomium, while in Phyllochetopterus it is incomplete dorsally
and shallow, allowing the prostomium to be seen easily. The
peristomium gives rise to a pair of tentacles which vary greatly |
in length, and in Phyllochetopterus there is also a second pair of
POLYCH-ETA FROM THE N.E. PACIFIC. 983
structures which are generally stated to be tentacles. They are
very small flat organs, lying on each side of the prostomium, and
generally covering the eyes. Claparéde showed, in the case of
P. socialis, that they contained two or three thin capillary sete.
This circumstance is easily verified in such species as I have
examined, and it leads me to suggest that we are in error 1n
regarding these structures as tentacles. They are much more
probably the reduced and modified notopodia of the peristomial
segment itself,
It may be of some advantage to dwell on this point at greater
length. In Annelids the peristomium 1s the segment behind the
prostomium, and it can be generally recognised and homologised
throughout the group. This fact is, however, due more to its
position than to any morphological characters which distinguish
it from those sueceeding, and it is generally recognised as the
most anterior of the trunk-segments, which has been considerably
modified in connection with the mouth. With regard to its
identity with the other segments of the trunk, Goodrich says :-—
“‘ Gareful modern researches (Vejdovsky, Wilson, etc.) have shown
that in Oligochetes the peristomium exhibits the essential charac-
ters of a true segment. It develops as a region surroun ding the
mouth, in which are formed a pair of mesoblastic somites which
become hollowed out to form the ccelom; a ganglionic thickening
is produced ventrally, which soon fuses with that of the succeeding
segment; a nephridium (head kidney) is developed. In the
Polychetes—in some cases, at all events,—it has been shown that
a pair of somites are formed in the peristomium, become hollowed
ont, and even give rise to peritoneal funnels. Nephridia are
almost invariably developed in this segment. In Polychetes,
moreover, a pair of lateral appendages are often developed, though
they generally become highly modified. In fact, it becomes
evident, when we examine the development and the adult structure
of the peristomium in the various groups of the Annelids, that it
is really a metamere strictly comparable to the posterior segments,
even when much modified owing to its position at the anterior
end of the animal.”
In the Syllids, the Polynoids, and other groups, the peristomium
consists of a segment which bears on each side a dorsal and
a ventral cirrus. That these are the remains of a typical para-
podium, in which the notopodium and neuropodium with their
setee have disappeared, is shown by a number of cases amongst
the Polynoids, which could probably be duplicated in other
families, where the first segment carries not only cirri but also
sete. In Polynoé extenuata, described by Claparéde, the peri-
stomium shows an aciculum and a couple of setae. In Pontogenia,
Sthenelais, Sigalion, and in Palmyra amongst the Palmyride, the
peristomium possesses a notopodium with several sete, and only
differs from the succeeding segments in the absence of neuropodial
sete.
These cases show that the peristomium may be an almost
984 . MR. F. A. POTTS ON
unmodified trunk-segment. On the other hand, it often occurs
that trunk-segments fuse with the peristomium and then lose
their parapodia, with the exception of the dorsal and ventral cirri.
So that the same process which has affected the peristomium may
modify the succeeding segments in a similar way.
The object of this digression is to show that the retention of a
parapodium in the peristomium of Phyllochetopterus is by no
means without parallel in other families of Polychets. The
constancy of the phenomenon makes it of generic value, and
suggests that possibly the rudimentary notopodium has under-
gone a change of function which we cannot yet appreciate.
DESCRIPTION OF A NEW Species, 7. anglica, From Brirish WATERS,
AND A CoMPARISON OF THOSE SPECIES OF THE GENUS WHICH
FORM ‘TRUE CoLONIEs.
PHYLLOCHEZTOPTERUS ANGLICA. (Pl. VI.; Text-figs. 9, 10, 12.)
Phyllochxtopterus of moderate size (2-12 em. in length), with
eye-spots. Anterior region with a variable number of segments
(13-16); @ single enlarged seta in each parapodium of the 4th
segment. Median region also with a variable number of segments
(11-25). Tubes creeping; often several run parallel to each
other, with short lateral connections. More than one individual in
the same system of tubes.
Locality. English Channel.
I obtained this species in March of the present year (1913),
while working at the laboratory of the Marine Biological Associa-
tion at Plymouth. Tubs of Chetopterus-tubes were brought in
from trawlers, apparently obtained from an area a little south of
the Eddystone. Attached to the Chetopterus-tubes were numbers
of other very slender tubes, often arranged in parallel bundles.
Sometimes they were embedded in the substance of the larger
tube, at other times they were entirely surrounded by large
colonies of Aleyoniwm, so that it seems that the communities of
Phyllochetopterus anglica are of comparatively long standing.
In the character of its colonies this species seems to be inter-
mediate between P. socialis and P. prolifica, and it will be
profitable to make a definite comparison :—
(1) In P. prolifica (Pl. IV.) the colony is usually contained in
a single, long, stolon-like tube (sometimes bifurcating) with a
number of very short branches communicating with the exterior,
and consists of a comparatively large number of smal] individuals
which reproduce asexually with rapidity (at least in the summer).
(2) In P. socialis the colony seems to occupy a large number
of parallel and adherent tubes, the communication between which
can only be made out with great difficulty. In each mass of ©
tubes the individuals are, however, of the same sex. A single
tube contains two or three individuals,
(3) In P. anglica (Pl. VI. figs. 15, 16) the colony is likewise
contained in a number of tubes, which tend to run parallel, but
are not,asarule,adherent. The open nature of the colony leaves
POLYCH.ETA FROM THE N.E, PACIFIC. 985
no but doubt that the tubes are connected. Usually each con-
stituent tube contains a single individual longer than those of
P. proiifica, Small subsidiary apertures may be placed at the end
of branchlets of the main tube, as in P. prolifica.
These distinctions which I have attempted to draw may, on
further examination of the genus, prove to be insufficiently
grounded. I feel quite certain, however, that P. prolifica in the
N.E. Pacific does not usually form the dense colonies charac-
teristic of P. socialis, nor do connected tubes run parallel as in
P. anglica.
It is possible, too, that these three species differ in the extent
to which asexual reproduction is developed in each. In particular,
it may be mentioned that P. prolifica is the only species in which
asexual reproduction was found to be proceeding at the moment
of discovery.* In P. anglica I was not successful in finding any
example which showed signs of recent regeneration. It is
probable that this phenomen takes place later in the year than
the time when my specimens were collected, but I incline te
beheve that autotomy and regeneration are never so frequent as
in P. prolifica, a cvcumstance which accounts for the smallei
number of individuals contained in much longer tubes.
In the morphological characters of the animals themselves, I
must confess to a considerable difficulty in distinguishing between
these three species. Myr. Crossland has pointed out that some
species of Phyllochcetopterus are easily recognised by definite
characters, like the number of inodified sete in the notopodium of
the fourth setigerous segment, the presence or absence of eye-
spots, and the number of sete in the notopodium of the
posterior region (C), as well as the character of the tubes. In all
four species, which have developed asexual reproduction, eye-spots
are present, a single strengthened seta is usually found in the
notopodium of the fourth segment and a single seta in each
notopodium of region C, while the tubes they inhabit are
creeping and branched. It seems to me that there is sufficient
difference between the forms here described to preserve them as
distinct species, though they must, from their morphological
character's, as well as from their manner of life and reproduction,
be classed as very nearly related. There are, moreover, differences
in the size, the number of segments in the several regions of the
body, and in the shape of the prostomium and peristomium
which help to supplement the biological peculiarities which I
have indicated above.
Comparison of the external Morphology of those Species of
Phyllochetopterus which form True Colonies.
(1) The Prostomium and Peristomium.—In P. anglica, the
prostomium (text-fig. 9) is rather broad, though small, and its
borders are marked by a line of dark pigment. The eyes ave
placed on the extreme side of the head, and are overlapped and
* This was in the summer (May-July).
986 MR. F. A. POLTS ON
hidden by the peristomial appendages, which are here small
and slender objects. The peristomium forms a well-developed
funnel. In P. prolifica, on, the other hand, though I have
examined a large number of individuals, I have experienced
considerable difficulty in making out the relations of the pro-
and peristomium. J think it is possible to say that the pro-
stomium is smaller, and less definite than in P. anglica, and that
the peristomial appendages are quite minute, and do not cover
the eyes. The peristomial funnel is complete.
I have been able to examine two of the species described by
Mr. Crossland, and preserved in the Museum of Zoology at
Cambridge, to test the variation which occurs in the different
species of Phyllochcetopterus. In P. elioti from Zanzibar the
pevistomial appendages are compa atively large and definite
structures, though they do not cover the eyes, the prostomium
is much better developed than in P. anglica aud P prolifica,
but the peristomial funnel completes little more than a semicircle
(Crossland, 2, pl. xiv. fig. 1).
Text-figure 9.
Phyllochetopterus anglica.
Dorsal view of head and anterior segments.
In P. pictus, also from the neighbourhood of Zanzibar, the
prostomium is rather large and fleshy, and the peristomium does
not form a funnel but a conical elevation, divided behind by a
median groove; the mouth is a small slit-like aperture. In
another species P. aciculigerus described by Crossland, the
peristomium is very much reduced, forming a cone with a rounded
mouth. The prostomium is small, but definite.
From the small series of species here examined, I have
ventured to draw some conclusions. The prostomium is always
a very definite organ, except in a species like P. prolifica, where
the peristomial collar is quite complete. The peristomium,
however, varies a good deal. In P. pictus and aciculigerus it 1s
small and rudimentary, while in other forms it is developed intoa
funnel—markedly incomplete, for instance, in P. elioti, less
so in P. anglica. With the completion of the peristomial
funnel in Mesochetopterus taylort and Chetopterus we have the
suppression of the prostomium and the complete disappearance
of the peristomial appendages. I think that the conical peri-
stomium is primitive and that the formation of the peristomial
funnel is a direct adaptation to microphagous habits.
POLYCHEHTA FROM THE N.E. PACIFIC. 987
(2) The Sete.—A good deal of time has been spent in
endeavouring to fix the value of the notopodial sete of the
anterior region as a basis of classification. It must, however, be
stated that the results of this enquiry are entirely negative. The
variation in the shape of the sete is almost endless, and seems to
occur indiscriminately in species and individual. The dorsalmost
sete in each parapodium are nearly always lanceolate and
symmetrical. Individual variations occur even here in the
Text-figure 10.
Phyllochetopterus anglica.
Series of notopodial sete from anterior region.
length and thickness of the head, and substantial modification in
the more ventral sete. The followi ing rough classification of types
of modification may be given as indicating the range of
variation :—
J. Shortening of the head, which remains symmetrical (text-
Tey Oh Age):
IT. An increasing asymmetry of the head.
(a) The head remains long and is drawn out into a
long asymmetrical tip (text- fig. LOR CSD):
(b) The head is much shortened, with a short tip and a
broad edge (text-fig. 10, E).
988 MR. F. A. POTTS ON
Minor variations are found, according to whether the sides of
the head are curved or straight, to the degree of attenuation and
curvature of the tip.
In none of the species which have been described hitherto
has a full examination of the notopodial sete been made. Such
figures and notes as are given are of little value, then, in fixing
the species; but they show, I think, that the same variations
which oceur in P. prolifica and anglica occur in all. I have been
able to compare these with the actual specimens of P. clapareden,
pictus, and elioti described by Crossland, and this examination
supports my conclusion that the notopodial sete are too variable
to base specific characters upon.
Text-figure 11.
Sek
We
A B ce
Phyllo chetopterus prolifica. Enlarged sete from 4th segeint.
The specially modified sete in the fourth segment are always
figured in descriptions of species, and their configuration is
sometimes considered to be of diagnostic value. There is, without
doubt, a recognisable type for each genus of the Chetopterids,
but the modifications of this are so many, varying even in the
same individual, that I find it impossible to regard the shape
of the seta as in any way a specific feature. In all species of
Phyllocheetopterus, the seta appears to end in a blunt elliptical
crown. One or both of the sides of this are raised into a cuspate
ridge, one ridge being generally higher than the other. The
number of cusps is highly variable, and so is their development.
Thus in P. prolifica (text-fig. 11, A, B) there are on the highest
side 3, 4, or 5 cusps, and generally the cusps have a fairly equal
development. In some cases, however, like text-fig. 11, C, one or
POLYCH ESTA FROM THE N.E. PACIFIC. 989
more of the cusps are of greatly inereased size. The lower side
of the crown is usually smooth or slightly crenulated.
In P. anglica the set are often very similar to those of
P. prolifica. Two sete are here figured to show the extent
of the variation in number and size of cusps. In one of them
(text-fig. 12, A) there are only two cusps, one of which is very
large. In the other (text-fig. 12, B) the appearance of a cuspate
lower border will be noticed.
Text-figure 12.
A B
Phyllochetopterus anglica.
Enlarged sete from 4th segment of two individuals,
The figure which Crossland has given of P. pictus shows an
enlarged seta of a type different from any which | have described
for P. prolifica and anglica. In text-fig. 138, C, however, IT have
drawn another seta from the 4th parapodium of a 2. pictus
collected by Crossland, and I think, it will be seen to be easily
derived from the seta shown in text-fig, 12, A (P. angliew).
The cusps on the lower border are better developed in the first,
but otherwise the two are strongly similar. ‘The bulging shaft
of the seta mentioned by Grossland is shown also in my
mounted specimen, but I ave observed similar phenomena in
P. prolifiea.
P. elioti is another species which from the published description
appears to have a very definite type of strengthened seta. 1
mounted two or three setz from Crossland’s specimen, and one of
these (text-fig. 13, B) agreed fairly well with his figure. On the
upper side the two external cusps are greatly enlarged, contrasting
with the two small intermediate cusps (not seen at all in Cross-
land’s figure), But another seta (text-fig. 13, A) showed an
oblique crown with small equal cusps, like the usual type of
P. prolifica ete.
It is very probable that the average seta of a species is different
990 MR. F A. POTTS ON
from that of another species, but the point I wish to emphasise
is that the sete of the fourth parapodium vary greatly, and a
statement as to their character can only be made after examination
of a large series of individuals. It has been shown in the
preceding part of this section that the other sete of the anterior
region are always variable, and it would have been rather
surprising if those modified setze which are found in the fourth
parapodium had been found to belong to types fixed for each
species.
Text-figure 13,
A, B. Phyllochetopterus elioti. C. P. pictus.
Enlarged sete from 4th segment.
Some Remarks ON THE GENERA Spicchetopterus AND
Phyllochetopterus.
Until 1856 Chetopterus was the only member of the family
known, but in that year Michael Sars (10 a) described ‘the
genus Spiochetopterus to include a species (S. typica) from
Norwegian waters, which differed from Chetopterus in possessing
long peristomial tentacles. In 1863, Grube (7) instituted a third
genus, Phyllochetopterus, for a worm from the Adriatic. But the
two forms are undoubtedly similar, and de Quatrefages, in his
‘Histoire des Annélés,’ goes so far as to include Grube’s polychet,
P. gracilis, in the earlier genus Spiochetopierus.
Spiochetopterus typica was described as living in a jointed
transparent tube. It has long peristomial tentacles, but a pair of
POLYCHETA FROM THE N.E. PACIFIC. 991
peristomial appendages was not observed. There are three well-
differentiated regions, the median of which contains two segments
with bifid foliaceous notopodia, exactly like those described in all
species of Phyllochetopterus. ‘There is only one strongly modified
seta in the 4th segment. In the posterior region there is a
bundle of fine sete in the notopodium, but the neuropodium is
stated to be without uncini.
Phyllochetopterus gracilis, as originally described by Grube,
‘possesses a pai of short peristomial tentacles. It is possible
that in the cases examined the greater part had been broken off,
bat Grube thinks this was unlikely. Or they may really be very
long peristomial appendages, the tentacles being entirely lost.
The observations are quite inadequate on this important pot.
The 4th segment of the anterior region appears to have more
than one strengthened seta on each side, though the number is
not actually given. There are two segments here also in the
median region, and they are similar in form to those of Spio-
chetopterus. Vhe posterior region contains segments, the noto-
podia of which each contains more than one seta; but Grube
expresses himself as uncertain whether the neuropodium
contains uncini or not.
Neither of these forms has ever been rediscovered so far
as I know, and so these descriptions remain still inadequate
and uncorrected. But as they stand, I share the opinion of
de Quatrefages concerning them, that no sufficient cause is shown
for placing the Adriatic form in a separate genus. The differ-
ences of the tentacles might be explained as errors of description,
and the variation of number of strengthened sete in the 4th
segment is unimportant. The presence or absence of uncini in
the neuropodium of the posterior segments is a moot point in
both, but it is probably their extremely small size which enabled
them to escape detection.
The next question which arises is whether these two forms are
similar to those better-known species which are grouped to-day
under the genus Phyllochetopterus. For that genus is charac-
terised by the possession of a pair of peristomial appendages, as well
as the long tentacles, and they may possibly have been overlooked
by Sars and Grube in their respective discoveries. ‘The numbers
of enlarged sete in the 4th segment and in the notopodium of
the posterior segments and that of the segments in the median
region are not definite generic characters. ‘The structure of the
segments of the median region of these two forms is identical
with the type usually associated with Phyllochetopterus. It is
however, I think, a matter of some importance that these two
early forms should be rediscovered and their position more
accurately defined. For the present, the generic name Phy/lo-
chetopterus must certainly be retained, and I trust it will not be
necessary to go back to the older genus Spiochwtopterus. But
in the table of the Cheetopterids which is given by Crossland he
includes Spiochetopterus as distinct from Phyllochetopterus through
992 MR. F. A. POTTS ON
its single pair of tentacles. I should like to point out that both
genera were originally described as possessing only a single pair
of peristomial processes, and that we are not ina position to
correct the description of the type-species.
LITERATURE.
(1) Chaparkpr, Epovarp.—Les Annélides Chétopodes du
Golfe de Naples. Seconde Partie. (Pp. 76-94 for
Cheetopterids. )
(2) Crosstanp, Crrit.—On the Marine Fauna of Zanzibar and
British East Africa, from Collections made by Cyril
Crossland in the Years 1901 and 1902.— Polycheeta.
Part I. Proc. Zool. Soe. London, 1908, vol. i. pp. 169-
176.
(3) Crosstanp, Cyrin.—The Polycheta of the Maldive
Archipelago, from the Collections made by J. Stanley
Gardiner in 1899. Proce. Zool. Soe. London, 1904,
vol. i. pp. 270-286, pls. xvili., xix. (For Phyllocheto-
pterus aciculigerus and gardinert.)
(4) Expers, Howarp.—A Study of the Life-History and Habits
of Cheetopter us variopedatus Renier et Cl: aparede. Journ.
Morph, Philadelphia, vol. xx. 1909, pp. 479-531,
pls. 1.111
(5) Gravier, C.—Annélides Polychétes de la Mer Rouge.
Nouv. Arch. Mus. d’Hist. Nat. sér. 4, t. viii, 1906. (For
Telepsavus bonhourei, see p. 191, pl. iii. figs. 209-213.)
(6) Gravier, C —Sur la régénération des extrémités du corps
chez le Chétopteére et chez la Marphyse sanguine, Bull.
Mus. Paris, 1909, pp. 14-17; Ann. Sci. Nat. (Zool.)
Paris, sér. 9, t. ix. 1909, pp. 129-155.
(7) Groupe, E.—Beschreibung neuer oder wenig bekannter
Anneliden. Arch. f. Naturgeschichte, 1863, p. 52.
(8) Ivanow, P.—Die Regeneration des vorderen und des
hinteren Korperendes bei Spirographis spallanzanit Viv.
Zeitschr, f. wiss, Zool. Bd. xci, pp. 511-558, Taf. 20-22.
(9) Larrurs, J. Joysux-.—Etude Monographique du Chéto-
ptére. Arch, Zool. Exp. Gén. sér. 2, t. vii. 1890,
pp. 245-360, pls. xvi.-xx.
(10) Lancrernans, P.—Ueber einige canarische Anneliden.
Nova Acta Leop. Car. Akad. Naturf. Bd. xlii. Nr. 3,
1881. (For Phyllochetopterus gracilis.)
(10a) Sars, M.—Fauna littoralis Norvegie, ii. 1856.
(11) Vaney, C, et Contr, A.— Recherches expérimentales sur la
régénération chez Spirographis spallanzanti (Viviani).
C. R. Soc. Biol. 1899, p. 973.
(12) Watson, A. T.—A Case of Regeneration in Polychet
ome Proc, Roy, Soc, ser, B, vol. Ixxvii. 1906,
pp. 332-6,
.
POLYCH.EYA FROM THE N.E. PACIFIC. 993
EXPLANATION OF THE PLATES.
Lettering.
a.s. Proximal and p.s. distal half of posterior segments.
cil.gr. Ciliated groove and enl. enlargements of the same to form accessory
feeding organs.
d. Dorsal lengthened sete of notopodium.
Jr. The frilled borders of the median region.
gl. Glandular epithelium of dorsal surface.
lab. The two ventral lips of the mouth.
neur. Neuropodium.
neur. The undivided neuropodium of the first segment of the median region.
neur.”, neur.® The double neuropodia of the second and third regions.
not. Notopodium.
not.', not.2, not. Notopodia of segments in median region.
per. Peristomial collar.
pl. Ventral plastron of anterior region.
pro. Prostomium.
st.t. Stumps of peristomial tentacle.
é. Peristomial tentacle.
w. Prolongation of the ventral plastron into median region.
Prate I.
Mesocetopterus taylori, sp. nu. Departure Bay, B.C.
Fig. 1. Dorsal view, comprising the anterior and median regions with the first
segments of the posterior region.
2. Lateral view of same to show dorsal approximation of the borders (/%.) of
the median segments, ete.
3. View of lead from above to show the prostomium completely surrounded
by the peristomium, and the slit-like mouth bordered by two ventral
lips.
(All three figures are drawn about twice the natural size.)
PrateE II.
Mesochetopterus minuta, sp.n. Cape Verde Is.
Fig. 4. Dorsal view showing the well-developed prostomium, continuous ciliated
groove, absence of frilled borders to median region, ete. X 20.
Puate III.
Mesochetopterus.
Fig.5. Ventral view of IZ. taylori, to show end of anterior and beginning of
median region. X 3.
. Lateral view of posterior region in WV. taylori. X 3.
. Lateral view otf end of median and beginning of posterior regions in
M. minuta. X 10.
. Hinder segments of posterior region in MW. minuta. X10.
. Anterior view of a parapodium in the anterior region of WZ. taylori.
Puate IV.
Phyllochetopterus prolifica, sp. nu.
Fig.10. A colony containing three adult worms, two of which (A, B) have just
separated off portions (A’, 3’) of the posterior region. The rounded
original end of the tube is seen at D. t
Departure Bay, British Columbia. Natural size.
11. A colony containing at least three adult worms. This shows two important
lateral branches and a large number of short branches bearing
apertures. That part of the system represented with close shading is the
older, the tube-walls being opaque, and that with distinct annulations is
newer, the transparent walls showing the worms within. (Two of these
wornis which alinost touch each other, lie in opposite directions.)
San Juan Archipelago, Washington. Natural size.
(In both figures, the anterior and median regions of the worm are shown dotted,
the posterior region very dark. The tentacles are omitted.)
Proc. Zoou. Soc.—1914, No. LX VII. 67
“I o>
۩ 00
994 ON POLYCH ATA FROM THE N.E, PACIFIC.
Prats V.
Phyllochetopterus prolifica.
A series of three individuals showing the stages in the regeneration of the anterior
region by a posterior fragment containing segments of the median and posterior
regions.
Fig. 12. Beginning of regeneration : anterior region represented by a tiny stump.
Sermentation is indicated by the superficial folds, and the peristomial
tentacles are seen as two apical outgrowths.
13. Anterior region is almost complete, but still very small: peristomial
tentacles short and fourth segment without enlarged sete.
14, Adult worm with fully formed anterior region. Tentacles completely
developed. The whole of the posterior region is not shown.
PratE VI.
Phyllochetopterus anglica, sp. i.
Fig.15. A colony containing two fully developed worms. It consists of two
parallel tubes with short connection. One tube shows the rounded
original end. Natural size.
16. A colony consisting of several parallel connected tubes, in part adherent to
each other. Only one worm was found inhabiting it. Natural size.
(In both figures, the anterior and median regions of the worm are shown dotted,
the posterior region black, The tentacles are shown. ‘The tubes where they
are transparent are shown unshaded, where opaque darkly shaded.)
1914 WATSON ela
Bale & Danielsson, Lt? imp.
BROOMIA PERPLEXA.
ON A NEW FOSSIL REPTILE. 995
. Broomia perpleva, gen. et sp. n., a Fossil Reptile from
South Africa. By D. M.S. Watson, M.Se., F.Z.8.,
Lecturer on Vertebrate Paleontology in University
College, London.
[Received June 3, 1914: Read November 24, 1914.]
(Plate VI.* and Text-figures 1-5.)
INDEX.
Page
Broomiaupenplerd. SeNeeb SPs le) qe... .ceiecsse-escaree cease 1002
Adelosaurus, gen. u....... SRE Mirae yb 1009
Description of Skull of Brains eaplicsa ee Soe eee 995
.; Wertebralicolumimnieente = eee ee ee OOS
Pe LULU Sipe ERR ee eee oe ssa asnn a eae ye 998
a Recthoralltoindl meena acca. secs ce ceaeeeeees 1 998
3 Boreylimippemrennsss-ccteces tock! cieesetaea ences, 1 1909
- Ilium and liana lisvals Peete eae, LOOO
Comparison with other groups of Reptiles ..................... 1003
Generaliconclusion smear ene ee ee OOD
Whilst collecting on the farm Hottentots Rivier, Field Cor-
netcy Gouph No. 3, District Beaufort West, Cape Province, from
the Pariasaurus zone, I was given a lump of the or diene
quartzitic sandstone of that horizon showing an extremely sharp
impression of a small lizard-like reptile. By careful development
I exposed a perfectly preserved car pus and tarsus with the actual
bone well preserved, and the specimen now allows of a very
complete account of the animal’s structure.
Skull—The skull is represented only by an exquisitely sharp
impression of the buccal surface of the palate, squeezes from
which show its structure with perfect clearness.
The basioccipital is not definitely visible, but it may be
represented by a rather faint impression behind the basisphenoid ;
it is, however, equally probable that this represents the atlas.
The basisphenoid is very broad, the lower surface is shallowly
concave, the lateral borders being raised into low, sharp-edged
ridges which shghtly separate posteriorly and end in ill-marked
tubera. Anteriorly the bone bears two well-marked basipterygoid
processes which, so far from projecting downwards, lie above the
general level of the lower surface of the bone; they are directed
forwards, and their flat articulating surfaces are nearly at right
angles to the length of the skull. Between these processes the
parasphenoid projects forward as a very narrow rostrum of
considerable length ; where it joins the basisphenoid it separates
distinct grooves on each side of the middle line which lead from
the palate into the skuJl. There is some evidence that the
parasphenoid terminated behind in a diamond-shaped expansion
on the lower surface of the basisphenoid.
There are no carotid foramina.
* For explanation of the Plate see p. 1010.
67*
996 IR. D. M. 8. WATSON ON
The pterygoid articulates by a pendunculate facet with the
basipterygoid process ; behind this the posterior ramus runs back
to the quadrate. It is shown in the specimen as a very narrow
strip, but in front there are appearances which suggest that the
part visible is really only a narrow rib on the lower surface of a
much broader bone, as in Captorhinus. In front the pterygoid is
not very clearly distinguishable from the palatine and ecto-
pterygoid. It bears three raised ridges, each covered with a
granulation of very small, closely-set teeth ; the posterior ridge is
Text-figure 1.
Broomia perplexa. Restoration of palate. 2.
B.Sp., Basisphenoid ; Jw., Jugal; Mer., Maxilla; Pal., Palatine; Par.Sp., Para-
sphenoid ; Pr.Jfv., Premaxilla; Pr.V., Prevomer; Pé., Pierygoid; Qu.,
Quadrate; Qu.Ju., Quadratojugal ; Tr, Transverse.
short, directed nearly laterally, and has only a single series of
teeth; the second ridge is rather longer, directed forward, and
bears two irregular rows of denticles; the third ridge forms the
inner margin ef the bone and runs forward as far as the specimen
allows it to be seen; this ridge is covered with a large number of
irregular rows of teeth, so that it forms a closely-set granular
area. ‘The surface of the bone between the dentigerous ridges is
depressed and between the two anterior ridges has a few smail
scattered teeth. There is a distinct process applied to the inner
A NEW FOSSIL REPTILE. QO7T
side of the lower jaw, not much depressed, and apparently formed
partly by the ectopterygoid, which bears no teeth. In this region
there are two small depressions which may be foramina.
A very remarkable feature is the very large size of the inter-
pterygoid vacuity ; it is probable that the pterygoids do not meet.
The palatine is a bone which bears a single dentigerous ridge
directly continuing the middle one on the pterygoid ; where the
two bones meet there is a small gap with no teeth; the anterior
end of the palatine seems to show a natural border at the back of
the internal nares.
The maxilla is not well shown, but in its anterior portion
bears a single row of very closely-set teeth which are wider from
side to side than they are long; posteriorly the teeth are small,
separated, and circular. The maxillary teeth seem to be shee
codont, and are very short and blunt, being directly apposed to
those of the lower jaw.
The quadrate is shown on the left side as a rather thick plate
curved lateral ly, so that its inner border comes into contact with
the front face of the posterior ramus of the pterygoid, and its
thin outer border forms part of the outer side of the skull.
Lying to the outside of the quadrate, but with its thin posterior
border within that bone, is seen the lower edge of a bone which is
very short antero-posteriorly. This bone can only be a quadrato-
jugal or a squamosal. Tightly applied to the outer surface of
this bone is the extreme tip of another, which might be a
squamosal if the other be a quadratojugal.
On the same side, lying in close relation to the posterior end of
the maxilla, is an ibe shaped bone which can only be the jugal; its
border all round seems to be a natural one. From its curvature
it is certain that the long limb cannot have reached back to the
quadratojugal, but formed the back of the orbit, which must
have been very large. As the squamosal and quadratojugal are
in their natural “position and the jugal is displaced, it seems
certain that there was no lower temporal arcade and that the
temporal region was cut away from below, as in lizards.
The lower jaw is in place and the left side of the palate is
perfectly preserved, so that there is no difficulty in making a
restoration of the palate. In such a restoration the pointed
shape and width of the skull are very noticeable, as are the
enormous interpterygoid vacuity and the fact that the articular
region of the quadrate lies far in advance of the basioccipital.
"The lower j jaw of the right side is perfectly preserved and fairly
well exposed. There is a small splenial entering the symphysis
and overlapping the angular behind ; the rather larger dentary
overlaps the outer side of the same bone. The angular is a large
boat-shaped bone forming the bottom of the jaw behind. ‘The
surangular and articular are not exposed, but on the left side a
short, high, very lizard-like coronoid process lies outside the
pterygo-transverse process.
The two rami are only loosely connected in front.
998 MR. D. M. S. WATSON ON
Vertebral coiumn.—The anterior vertebre are not known,
having been in the other side of the block, which was not
recovered. The whole skeleton lies in position, so that from
measurements it is possible to obtain the length of the missing
part; if, as seems most probable, the anterior vertebrae were of
the same length as those behind them, eight are missing, giving
a total of twenty-four presacrals. All the presacrals preserved
are much alike in form and size. The centrum is small, with a
wide hourglass-shaped notochordal canal running through it; it
is expanded at the articular ends and somewhat constricted in
the middle ; the lower surface is rounded but slightly flattened.
The arches are very wide and the neural canal enormous, much
wider than it is high. The last presacral gives satisfactory
evidence that the zygapophysial articulating surfaces were flat
and placed horizontally; the anterior zygapophyses only project
very slightly, if at all, in front of the centrum. The transverse
process best seen in the 12th and 13th presacrals is short, and
extends from a point on the arch near or on the neurocentral
suture up to the process which supports the prezygapophysis, but
the articular facet for the rib begins some distance behind its
anterior end. There are intercentra throughout the column
except between the sacrals. There are two sacrals of the same
length as the presacrals; their centra are, however, more robust,
and have a very pronounced carination of the lower surface.
The sacral ribs are largely attached to the centra.
The first five caudal vertebrae, which alone are preserved, are of
the same length as the sacrals, but the centra are constricted and
rapidly lose the carination of the sacrals: there are apparently
intercentra between the first and second and all succeeding caudals,
but at what point these take the character of hemal arches is
not shown, although that between the 4th and 5th is a chevron
bone. ;
Ribs.—The presacral ribs are all single-headed, holospondylous ;
their articular end is somewhat swollen but is not very broad, as
it is in Dicynodon for example. The ribs are long, considerably
curved, quite slender, and ribbed in front.
The anterior sacral rib is short and strong; it has a large flat
surface for the ilium, and arises from the conjoint centrum and
neural arch rather far forward. The posterior rib is longer, but
perhaps not so strong as the anterior; it has a very large
articulation with the ilium, and its distal end is in contact with
that of the first sacral rib.
The caudal ribs of the first four vertebre are long simple
processes fused on to the body of the vertebre.
There are faint traces of abdominal ribs.
Pectoral Girdle.—The pectoral girdle is in position, and so far as
it is shown very well preserved. ;
The interclavicle is a large bone with a rhomboidal head produced
at the lateral angles into short processes, and with a very long
narrow stem. The head has its front edges bevelled off and
A NEW FOSSIL REPTILE. 999
recessed for the clavicles, and the under surface of the stem is
also recessed, apparently for the inner borders of the coracoids.
This implies that the interclavicle largely lay above the coracoids,
a feature only paralleled by the Plesiosaurs. The evidence for
this curious arrangement is very much str engthened by the fact
that the stem is broken, and the lower part with the right
coracoid underlying it is pressed up, whilst the left coracoid
retains its natural position in relation to the anterior end of the
interclavicle.
The lower end of the left clavicle is well preserved; it is rather
wide and thick, lies along the anterior border of the head of the
interclavicle, and shows a very feeble sculpturing of pits and
grooves. The coracoid and scapula of each side are fused
together, and only the lower part of the joint-bone is exposed.
The coracoid is a large flattish bone with curved borders; it bears
a strong process which carries the lower and posterior part
of the glenoid cavity, behind which the bone is continued for some
distance. There is a small, oval, coracoid foramen in the groove
which continues the glenoid cavity forward. There is a powerful
rounded supraglenoid process borne by the scapula, and some
slight evidence of a glenoid foramen. The right scapulo-coracoid
shows that the whole bone forms about a quadrant of a circle.
There is no reason to suppose that more than a single coracoidal
element was present: this being no doubt, as Williston believes,
the anterior of the two of the Cotylosaur shoulder-girdle.
Fore limb.—The upper part of the right humerus is shown
from below; the left humerus is badly exposed, but shows the
length of the bone and something of its distal end. The bone is
very slender; it is slightly expanded at both ends, and no doubt
somewhat twisted. The head is not well exposed; there is a
short but relatively powerful radial crest which rather rapidly
subsides on to the shaft. Of the distal end, all that can be said
is that it is exceptionally well ossified and finished, with a round
condyle for the radius, facing at right angles to the shaft, and a
facet for the sigmoidal fossa of the ulna at the end.
The radius is a long slender bone slightly expanded at the
ends. The ulna is a slender bone, with the upper end thickened
and produced into a very pronounced olecranon process.
Although the carpus is only a centimetre square its structure
is shown with diagrammatic clearness on the right side, where
it is exposed on its palmar aspect and has the actual bone well
preserved.
There are three large proximal carpals, anda slight suggestion
of a small pisiform lying a little removed on the ulnar side. There
are three centralia which completely separate the proximal and
distal rows of the carpus. The most ulnar of these is very small,
articulates with the ulnare, median centrale, and fourth distal
carpal. The median centrale is one of the largest bones of the
wrist, articulates with the ulnare, intermedium, radiale, radial
centrale, third and fourth distal carpals, and the ulnar centrale.
1000 MR. D. M. 8S. WATSON ON
The radial centrale is a large bone articulating with the radiale,
median centrale, and the first three distal carpalia. There are
five distal carpals, all except the fifth being large bones in
mutual contact. The fifth isa smailer bone, forming only part
of the support of the fifth metacarpal. The carpus as a whole is
remarkable for its thorough ossification and the accuracy of fit of
its elements,
Text-figure 2.
Broomia perplexa.
Outline drawing of right carpus and manus as preserved.
R. Radius, U. Ula. XX 2 approx.
Niwm.—Neither ilium is quite complete, but the two supple-
ment one another so as to give a good idea of the whole bone,
There is no preacetabular projection, the blade of the bone ex-
tending upwards and backwards from the strong process which
projects over the acetabulum. The outer surface of the bone
has a strong ridge running horizontally across it, and the upper
end shows a faint grooving for muscle-insertion.
The pubis and ischium cannot be exposed.
The left femur is only shown in section, but the right gives
some idea of the form of the bone. It is slender and sigmoidally
curved. The articular facet at the distal end is well rounded ;
the upper end of the bone is flattened, but bears a strong
trochanteric ridge. The tibia and fibula are very slender bones
shown only in section. The left tarsus is beautifully shown; it
has the bone preserved and is exposed on its dorsal surface. Its
A NEW FOSSIL REPTILE. 1001
proximal row consists of two very large flat bones which meet in
the middle ina long articulation broken by notches which to-
gether form a small foramen. One of these bones is the fibulare,
the other in all probability the fused tibiale and intermedium.
The distal row consists of five bones, of which the fourth is very
large and articulates with the two proximal tarsals. The first,
second, and third distal tarsals are separated from the tibiale
intermedium by two centralia which form a median row in the
tarsus.
Text-figure 3.
Ske:
OS
NEN
NEN
CEN
x
.
N
]
ee
gd.
i
|
Broomia perplexa.
Outline drawing of the left hind leg as preserved. X 1.
The first metatarsal is not exposed; the base of the second is
seen supported by its tarsal. The third is a slender bone of
considerable length. The fourth is even longer, being more than
half as Jong as the fibula. The fifth metatarsal is short, only
slightly more than half as long as the fourth. In the specimen
all the metatarsals lie parallel and close up to one another.
This little animal, as restored in text-fig. 4, is thoroughly
lizard-like in build and obviously led a lizard-like lite on perfectly
dry land. It may perhaps, as suggested by its large claws and
very slender limb, have been to some extent arboreal.
The little lizard-like animal, almost the whole of whose
structure is described in the foregoing account, is obviously
distinct from any known South African form, and as it is the most
striking new form which I collected in that country, I propose
1002 MR, D. M. S. WATSON ON
for it the name of Broomia perplexa, gen. et sp. 0., in token of
my admiration of Dr. R. Broom’s work on early Tetrapods. In
Text-figure 4,
Broomia perplexa.
Restoration of the skeleton from the ventral aspect with the abdominal
ribs omitted. X +.
discussing its systematic position it will, I think, be most con-
venient to compare it in detail with all the great groups of
A NEW FOSSIL REPTILE. 1003
early reptiles which are well enough known to make a com-
parison of much value, and then to discuss some of the less known
Permian types which resemble it.
Text-figure 5.
Adelosaurus huxleyi (Hancock & Howse).
Left fore limb and right scapulo-coracoid. X 15.
From the type-specimen in the Hancock Museum, Newcastle-on-Tyne.
Comparison with Cotylosaurs.
The palate with its abundant armature of teeth is somewhat
similar to that of the Captorhinide, in which the distribution of
the teeth and even some details, such as the shape of the ptery-
goids and the length of the parasphenoid, are identical.
The basisphenoid of Broomia differs, however, from that of all
known Cotylosaurs in its very great breadth and only very
slightly marked tubera. i, as seems probable, the side wall
of the skull was cut away in Broomia, we have a very striking
4
1004 MR. D. M. 8S. WATSON ON
'
difference, but one which the example of the Chelonia teaches us
might have occurred very rapidly.
The vertebral column of Broomia is very similar to that of the
Captorhinids, having small completely perforated centra, heavy
neural arches and horizontal zygapophysial articulations, and
small intercentra. The rib articulations, however, are smaller in
our type.
The limb-girdles of Broomia are totally unlike those of any
Cotylosaur in their great slenderness. The shoulder-girdle differs
very markedly from that of the Captorhinide, but faintly recalls
that of Seymouria in its peculiar interclavicle with a diamond-
shaped head and in the clavicle with its curious expanded lower
end. The loss of the posterior coracoidal element is paralleled
by Seymouria, but in that type if I interpret Prof. Williston’s
description accurately, the real Coracoid was present although it
was not ossified and contributed to the glenoid cavity, which, as
in the majority of Carboniferous reptiles, had the peculiar screw-
shaped form most typically shown in Diadectes and Hryops.
Traces of this former possession of this type of glenoid cavity are
to be seen in Broomia, but the conditions there are different, in
that the whole glenoid cavity is carried by the scapula and the
single coracoidal element, which extends backward for some
distance behind it.
The humerus is distinguished trom that of any Cotylosaur by
its slenderness, but is probably structurally similar to that of
Captorhinus.
The carpus of Broomia is unique, no other form being known
in which the centralia completely separate the proximal and
distal rows of carpals.
The ilium, which alone of the pelvic bones is known in Broomia,
differs from that of all Cotylosaurs in its slenderness and its
sloping anterior border.
The femur is so badly exposed that it is difficult to compare it
with that of Cotylosaurs, from which, however, it differs in its
extreme slenderness. The tarsus differs from that of any known
Cotylosaur in the presence of two centralia.
These resemblances, particularly those in the vertebral column,
seem to show that Broomia has descended from some Cotylosaur; ,
the differences, lying chiefly in the build and limb-skeleton, are in
general advances of an adaptive nature, but the tarsus is the
most primitive known amongst reptiles, and the carpus cannot at.
present be explained.
Comparison with the Therapsid stock.
Varanosaurus and its immediate allies amon gst the Poliosauridz
are the most primitive known members of the Therapsid line,
and as they are comparatively small and lightly built reptiles,
offer an exceptionally favourable field for comparison with
Broomia.
A NEW FOSSIL REPTILE. 1005
The palate of Broomia is not very unlike that of Varanosaurus,
but differs in the wide basisphenoid, the very large inter-
pterygoidal vacuity, and the more abundant teeth. ‘The lower
jaw of Broomia is quite distinct from that of any Therapsid in
that it has a boat-shaped angular in place of the characteristic
flat-notched angular of the Therapsid. The vertebral column is
also somewhat similar to that of a Poliosaurian, but the neural
arches are heavier and the rib articulations not so wide. The
shoulder-girdle differs in that the shoulder-girdle of the Therapsids
always has a posterior coracoidal element which may be only
cartilaginous but contributes to the glenoid cavity.
The dium of Lroomia is strikingly like that of Pacilospen-
dylus, and not unlike that of Varanosaurus. The foot resembles
in some ways that of Varanosaywrus and is almost identical
with that of Ophiacodon, which P.iof. Williston believes to belong
to that group.
The difference in the lower jaws shows at once that Broomia
does not belong to the Therapsid line, and the resemblances
between Varanosaurus and Ophiacodon and Broomia seem to be
in general either primitive features or adaptive ones.
ft have compared roomia with Casea but see no special
resemblance between them. (Casea is perhaps an extremely
early offshoot from the Therapsid stock before it had acquired its
characteristic angular.)
Broonia has an obvious superficial resemblance to Arwoscelis
in that they are both very lightly built reptiles of small size. It
is at present difficult to compare them in detail. From the
published accounts of W illiston I have been able to find the
following resemblances :—
In both there are teeth on all the bones of the palate. If
Broom’s Ophiodewus, founded on the specimens of ‘ Bolosawrus”
figured by Case, is really Arwoscelis, then there 1s a very striking
sunilarity in the palate of the two types, in the large inter-
pterygoidal space, very long parasphenoid, and general structure.
The dorsal vertebre are similar in their slender notochordal
centra and heavy arches. The ribs are similar in having only a
single slightly expanded head which articulates with the arch
and centrum near the front end of the vertebra throughout the
series. The sacrum of dArwoscelis is said to be almost indistin-
guishable from that of lizards; that of Lroomia also resembles the
same forms. The tail is long in both types.
IT can find no characters in which the incompletely known
humerus of Broomia differs from that of Arwoscelis. The femur
and tibia of the two types seem to agree.
The more important known differences between the two types
are that Arwoscelis retains the primitive two coracoidal elements
and that there is no trace in that type of the cut-away side of
the skull of Broomia. At the same time it must be remembered
that the facts are not certainly known in our fossil and that the
jugals of the two forms have a considerable resemblance.
1006 MR. D. M. S. WATSON ON
On the whole, there is nothing in the known structures of
these two animals to prohibit a fairly close resemblance between
them, but until Prof. Williston’s full description is published it
is impossible to go beyond this.
By far the most interesting comparison is between Broomia
and a lizard.
It is certain that the lizard palate must have been derived
from one generally resembling that of Lroomia, and it is probable
that it may have specially resembled that type in the possession
of a very large interpterygoid vacuity and a very large para-
sphenoid.
The basisphenoid of Groomia at once recalls that of a lizard,
but I know of none that really resembles it.
The lower jaw of Lroomia is sutticiently generalised to have
given rise to that of lizards, and very characteristic of that
group is the short upstanding coronoid process.
If the side of the temporal region of the skull be really cut
away in Lroomia, we have a very striking resemblance to the
Lizard type, where the narrowing of the primitiv ely single arch
has produced the well-known present-day structure.
The geckos have notochordal centra and intercentra, as has
Lroomia.
The articulation of the single-headed rib of Broomia is
essentially similar to that of a lizard. The sacrum is also
sunilar in the two groups of reptiles.
The pectoral girdle of Broomia is extraordinarily similar to
that which the primitive lizards must have possessed in the
following features :—
The reduction of the coracoidal elements to one on each side:
this being, as Prof. Williston has pointed out, the anterior of the
two of primitive reptiles.
The long slender interclavicle with a rhomboidal head is a type
from which the characteristic cross-shaped interclavicle of a lizard
could be derived. A T-shaped interclavicle could not have
produced this form.
The somewhat expanded lower end of the clayicle is also a
feature which was apparently present in the early lizards.
There is nothing specially characteristic about the bones of the
fore-leg in lizards, and they could be derived from those of
Lroomia.
The carpus of Lroomia differs as much from that of any lizard
as it does from that of all other reptiles.
The ium of Sroomia is completely lizard-like in its antero-
ventral slope. The hind leg of Lroomia is not specially lizard-
like.
The only feature which we would expect to be present in an
ancestral lizard which does not occur in Broomia is that modi-
fication of the fifth digit, perhaps a divarication, which led to the
modified fifth metatarsal found in Lizards, Sphenodon, Chelonia,
Thecodonts, Crocodilia, ete.
A NEW FOSSIL REPTILE. 1007
The skeleton of Sphenodon is so thoroughly lizard-like that
Broomia resembles it nearly as much as it does a lizard, but
there is pretty clear evidence that there was not a lower temporal
areade like that of Sphenodon.
The only other group with which it seems necessary to compare
Lroomia is the Mesosauria. The skull is unknown in this type.
The vertebre differ in their much more massive arches, and mm
the mode of articulation of the ribs. The pectoral girdles of the
two types are generally similar, but the interclavicle of Jeso-
saurus is T-shaped. The carpus differs by the absence of
centralia in Mesosaurus. The pelvis differs in the shape of the
ihum. The tarsus differs in the complete loss of centralia in
Mesosaurus. It thus seems certain that the two types have little
to do one with the other.
Only two, Heliosaurus and Heliophilus, of the little-known types
from South Africa agree at all with Broomia.
Lreomia resembles them in the following features :-—
1. The sharply pointed but relatively short skull.
2. The shape of the pterygoid in Heliosaurus.
3. The position of the quadrate in advance of the basi-
occipital condyle in Heliosaurus.
4. The presence of a distinct neck.
5. The presence of intercentra throughout the vertebral
column.
. The heavy neural arches.
The single-headed ribs.
The similar number of presacral vertebra.
. The shoulder-girdle of Heliosauwrus much resembles that
of Broomia.
10. The slender limbs.
These resemblances, although thev are to some extent due to
the retention of primitive features, do seem to show that there is
some real connection between the three animals. In /eliosawrus,
however, as Broom has shown, there is some evidence of the
presence of a quadratojugal arcade, which is apparently lacking
in Broomia. When I examined the type-specimen of Heliosawrus
some time ago I was not specially interested in it, but even at
that time thought it conceivable that the apparent lower arcade
might be the upper edge of the lower jaw. Heliosawrus is of
interest because of the presence over its dorsal region of small
bony scutes identical with the osteoderms of lizards.
Of the European forms, Aphelosaurus and Kadaliosaurus from
the Lower Permian, and Proterosaurus and the animal known as
Proterosaurus huxleyi from the Upper Permian, present some
resemblance to Broomia in that they are slender lizard-like
reptiles. Kadaliosawrus from the Rothliegende of Dresden is
regarded by Prof. Williston, who has examined its remains, as
being extremely similar to Arwoscelis : in fact, he stated that there
1008 MR. D. M. S. WATSON ON
ave no visible differences between them; it need not, therefore, be
further considered.
Apheloswurus is unfortunately very little known, but its
shoulder-girdle seems very similar to that of Broomia in the large
size of the coracoidal part. The slender limbs, however, differ in
the fact that they are all of much the same length, and also in
the much greater narrowness and compactness of the tarsus.
The animal is, in fact, so incompletely known that little can be
said about it.
Proterosaurus is a very interesting form which is, however, still
very little known.
The skull, as known from the single, very imperfect example in
the College of Surgeons Museum, is pointed and has teeth on
the palate. There is apparently no evidence of the presence
of an upper temporal vacuity, and nothing can be said of the
condition of the temporal region. It differs from Broomia in its
very long neck and in the enlarged cervicals.
The vertebre differ in their light neural arches, but the rib
articulation is essentially similar in the two types. The sacrum
is fairly well preserved in the Newcastle Museum specimen from
Fulwell, Durham; it is composed of two vertebre carrying large
sacral ribs which resemble extremely those of Pacilospondylus as
figured by Case, and to a less extent those of Broomia.
The shoulder-girdle has a very large, presumably single, cora-
coidal element, and a rather slender scapula, ‘The large inter-
clavicle, with an expanded upper end, is not altogether unlike
that of Broomia.
The limbs are considerably more massive than those of Broomia
and are still very imperfectly known, and, as in that type, the
hind limbs are considerably larger than the fore. ;
On our present knowledge of Proterosaurus it is impossible to
be.certain of its systematic position, but it appears not improbable
that it has something to do with Lroomia.
The animal described by Howse and Hancock as Proterosaurus
hucleyi iz quite distinct generically from Proterosaurus and may
have no real connection with that animal.
It is a small form with a long neck in which, however, the
cervicals are not elongated. The centra are large and biconcave
and the arches heavy. The ribs are single-headed. There are
apparently intercentra present. The shoulder-girdle is fairly
well shown. The scapula is a bone with no special features, and
the single coracoidal element is very large and singularly lizard-
like. The clavicle has an expanded lower end. The limb-bones
are incompletely ossified.
The humerus is remarkable for the very slight expansion of its
extremities and the absence of a definite crest.
There is a small entepicondylar foramen.
The radius and ulna are small bones with no particular
characters.
The left carpus is perfectly preserved.
A NEW FOSSIL REPTILE. 1009
There are four large proximal carpals and two well-ossified
centralia, the radial e which forms part of the border of the
carpus.
There are only three (distal carpals, the first, third, and fourth,
but there is an obvious space for the second, which was either
cartilaginous or has dropped out. There is certainly no fifth
carpal, The metacarpals and phalanges are relatively rather
massive and taper rather rapidly. It is possible that the fifth
had only two phalanges.
The ilium is a small bone, only the inner aspect of which is
known but whose outline is like that of Belodon.
The affinities of this animal, for which the new genus
Adelosaurus may be founded, are quite obseure; the fore limb is
not very unlike that of Sphenodon i in some featur es, and it is not
improbable that the type may be connected with an Archisaurian
stoek.
The preceding discussion will, I think, have shown that
Broomia cannot be placed in any of the well-known Orders of
reptiles which occur in the Permian rocks of the world. It seems
not improbable that it is connected in some way with the earlier
Areoscelis and with the later lizards, but the absence of all
knowledge of the temporal region of the skull and what is
probably still more important, of the neural cranium, makes this -
resemblance rest on a very insecure foundation,
Comparison with other slender-limbed Permian forms, so far
as is possible from the very imperfect material available, shows
that whilst there are certain general resemblances between them
there are also many important differences which make it very
inadvisable to definitely group them together.
The fact that, whilst we know Almost the whole structure of
Broomia, we are incapable of doing more than guess, at its
affinities, owing to the absence of knowledge of the upper part
of the skull and of the brain-case, shows how very few are the
characters on which we really rely in estimating the affinities of
a reptile.
I am indebted to the Percy Sladen Trustees for assistance in
visiting South Africa, and especially to G. Gordon, Esq., of
Hottentots Rivier, to whose interest and hospitality I owe, not
only the beautiful skeleton of Broomia, but also many other fine
specimens. I have to thank the authorities of the Northumber-
Jand and Durham Natural History Society and Mr. E. L. Gill,
the Director of the Hancock Museum, for permission to examine
the type-specimen of ‘‘ Proterosaurus hualeyi.”
Finally, I wish to thank Mr. Pittock, of University College, for
the excellent photographs from squeezes, and Mr. H. E. Herring
for the photograph of the block.
Proc. Zoo. Soc.—1914, No. LX. VIII. 68
1010
eS
=
mow bh
oo
ON A NEW FOSSIL REPTILE.
EXPLANATION OF PLATE VI.
Broomia perplexa, gen, et sp. n.
. Untouched photograph of a squeeze of the palate of the type-specimen. X 1}.
In this figure the matrix was covered with a thin wash of white before the
photograph was taken.
. Pectoral girdle. Photograph as in fig. 1. X 13.
. Middle dorsal vertebrie. Photograph as in fig. 1. X14.
. Sacrum. Photograph as in fig. 1. x14.
. Left ilium. Photograph as in fig. 1. 1%.
1
4
Photograph of the actual’ specimen. X 1. Showing the right hind leg,
posterior pre-sacral vertebrae, the last three showing clearly the immense
size of the neural canal, the sacrum, and the caudal vertebrae.
PZ. Ss, WA WATS ONG aval
- =~
Bale & Danielsson, Lt° imp. .
EUNOTOSAURUS AFRICANUS.
MR. D. M. 8S. WATSON ON THE CHELONTA. 1011
93. Hunotosaurus africanus Seeley, and the Ancestry of
the Chelonia. By D. M. 8. Watson, M.Sc., F.Z.S.,
Lecturer on Vertebrate Paleontology in University
College, London.
[Received June 3, 1914: Read November 24, 1914.]
(Plate VII.* & Text-figure 1.)
INDEX.
Page
Origin and Structure of the Chelomia ........................... 1011
Description of Skeleton of Hunotosaurus ..................... 1015
Relationship of Hunotosaurus to the Chelonia ............... 1019
The great group of the Chelonia is of unusual interest, because
it is the only example of a persistent and world-wide Order whose
structure is entirely dependent on a bizarre specialization : the
development of the shell.
Tortoises are first known from the Upper Trias of Germany,
where they are represented by extremely typical and apparently
not markedly primitive forms.
By an analysis of the structures common to all Chelonia, it
is possible to form some idea of the chief features which must
have been present in the ancestors of the group. Although
it is impossible to determine whether or not these characters
were actually all present together in any type, it will be con-
venient to add them together to form an imaginary animal
which may be called ‘ Archichelone.”
Skull_—The skull of the Chelonia has been discussed by many
authors, most recently by Hay, whose conclusion is adopted: in
this paper.
Since the description of Sphenodon by Dr. Giinther, it has been
repeatedly pointed out that, in such a type as Chelone with a
completely roofed skull, the four bones—post-orbital, squamosal,
jugal, and quadratojugal—which form the lateral temporal arcades
of Sphenodon are present. If I do not misinterpret their opinions,
most of those who have discussed the subject believe that the
skull of Chelone has actually arisen from a two-arched form by a
secondary enlargement of these bones, until they unite by their
edges. This conclusion is rendered improbable by the fact that
some Cotylosaurs, Labidosaurus and Pariasaurus for example,
whose skulls have certainly always been completely roofed, have
an identical arrangement of bones in the temporal region.
' In many Chelonia, apparently without any correlation with the
animal’s habits other than the power of retracting the neck, the
temporal region is open; but this opening never results in
the formation of a two-arched form—in all-cases it has been
* Por explanation of the Plate see p. 1020.
68*
1012 MR. D. M. S. WATSON ON
arrived at by the eating away of a solid cranial roof either from
the back or from below. ‘The former process results in the loss
of a parieto-squamosal connection and the retention of that
between the jugal and quadratojugal, asin Trionyx: the latter
produces a skull like that of Chelys, in which the squamoso-
parietal connection alone persists. Extension of either process
results in the complete loss of both connections, as in Cistaudo.
‘The only view which really fits the case is that the primitive
Chelonian skuil was completely roofed like that of Chelone or
a Cotylosaurian reptile. This view is supported by the pre-
ponderance of roofed skulls in early fossil forms.
It is not at all difficult to derive the Chelonian skull from that of
a Cotylosaurian reptile. The chief differences in the palate, the
total loss of teeth, the reduced pterygoids and their firm union
with the basis cranii, and the rudimentary secondary palate, are
apparently dependent on the gradual development, of the horny
beak. The unrelated group of the Anomodontia (Dicynodontia)
show the gradual extension of a horny beak, followed by the
total loss of teeth, the reduction of the pterygoid and its close
union with the basisphenoid, and the development of a small
secondary palate which in such features as the fusion of the
prevomeys is extremely like that of a Chelonian.
Many of the peculiar features of the posterior part of the
Chelonian skull are due to the necessity of adequately supporting
the very powerful vertically placed quadrate.
Other remarkable features in which all Chelonians agree are the
loss of the postfrontal and prefrontal (or lachrymal), and the
usual absence of nasals ; with the exception of the last, these losses.
have repeatedly occurred amongst reptiles. A much more
striking feature is the loss of the facial processes of the pre-
maxillee, so that the bony external nares areunited. ‘This change
is only paralleled in the Mammalia, of which group it is one of
the most striking peculiarities.
‘“‘ Archichelone,” then, had a roofed skull with a primitive
reptilian palate, teeth, probably only a squamosal of the temporal
elements [and no facial processes of the premaxille].
WVeck.—Cope pointed out that all tortoises could be divided
into two classes according to the mode of retracting the neck,
either vertically in the Cryptodeira (to which may be added the
Trionycids and Dermochelys) or laterally in the Pleurodeira. This
feature to all appearances affords a foundation for an absolute
division of all known types, a fact the meaning of which is plain.
‘* Archichelone” must have had a long and flexible neck, capable
of bending in all directions ; and after the development of the
shell, w hen it was desirable to withdraw the head for protection,
the two lines gradually became distinct, any intermediate con-
ditions being obviously mechanically undesirable.
, aeAsvall known Chelonia have eight cervical vertebra we are
eae in claiming that number for “ Archichelone.”
discussion of the trunk I shall aigsoaen
THE CHELONIA, 1013
Dermochelys, as the evidence now seems conclusive that that type
has originated in Tertiary times from a Cryptodeiran ancestor.
Inall Chelonia the trunk is very markedly separated from the
neck and tail, being very broad and completely surrounded by
the shell. The shell of all Chelonia in which it is not reduced
has the following structure :—
There is a median dorsal row of dermal plates. There is
always a nuchal, sometimes a postnuchal, then eight or fewer
neurals and one or two pygals. The whole of this series seems to
be homologous, but the eight neurals are distinguished from the
other meen Glemients by their definite enmalesion with the
neural spines of the dorsal vertebra. It seems certain that at
one time the nuchal ete. were also directly associated with the
anterior and posterior dorsal and sacral vertebra, and that they
lost this primitive connection after the formation of the shell,
either by its exaggerated growth or by a shortening of these
regions of the vertebral column, so as to allow of the retraction
of the head and limbs within the shell.
All Chelonia have a row of costal plates on each side of the
neurals ; these are invariably eight in number and are fused with
the dorsal ribs from the second to the ninth. The development
of these plates shows that they are of purely dermal origin, and
they may be directly compared with the paired scutes found in
Crocodiles and many allied reptiles.
The fact that they have only fused with eight of the ten dorsal
vertebre shows that these segments in ‘‘ Archichelone ” were in
some way specialised and distinguished from the first and last
dorsals.
The marginal elements of the shell are not correlated with the
ribs, and perhaps represent a different order of scutes as is
suggested by Versluys.
The plastron of Cryptodeires and some Pleurodeires consists of
a single median and four pairs of plates. In most Pleurodeires
and Amphichelydia an extra pair of plates is inserted, and in the
remarkable Triassic Protochersis recently described by Fraas
two extra pairs.
The median and the anterior pair of plastrals are universally
recognised as the interclavicle and clavicles. The other pairs are
usually homologised with ventral ribs: that they resemble the
latter in being ventral dermal ossifications is of course true, but
I do not think it at all follows that they are necessarily derived
from them.
There is, for example, never any trace of a median or other
lateral rows in the plastron of Chelonia, whilst abdominal ribs
usually have an angulated median and more than one lateral row
of elements.
Furthermore, as “ Archichelone ” certainly had a series of dorsal
scutes comparable to those of a crocodile, it is by no means
unlikely that it resembled many members of that group in having g
ventral scutes not homologous with abdominal ribs.
1014 MR. D. M. S. WATSON ON
The fact that in early times tortoises with mesoplastra were pro-
portionately commoner and more widely spread than to-day, and
that Triassic forms may have two pairs of mesoplastra, suggests
that the number in still earlier forms was even greater, and possibly
- derived from a condition in “ Archichelone” where there was a
pair of plastrals to each segment of the middle dorsal region.
The fact that adaptation to aquatic life almost always implies
degeneration of the shell in Chelonia, shows conclusively that
“« Archichelone”’ must have been a land form.
Limb-girdles.—The most striking feature of the limb-girdles of
Chelonia is that they lie within the shell and ribs. It is plain
that this condition was gradually acquired after the development
of the shell by the migration of the pectoral girdle backwards, and
that further growth of the front and back of the shell which we
have already found necessary to account for the dissociation of
the nuchal and pygal elements from the vertebrae to which they
probably originally belonged. The whole arrangement is designed
to allow of the retraction of the limbs for protection.
~ In “ Archichelone ” the pectoral girdle was undoubtedly in the
usual position overlying the first dorsal ribs, and must have been
narrow relatively to the bulk of the trunk to allow of its passage
back within the ribs.
The pelvic girdle must also have been very narrow.
This shows that the neck and tail of ‘‘ Archichelone” must have
been sharply marked off from the trunk by their smaller diameter.
It is impossible to discover the structure of the limb-girdles of
primitive Chelonia from those we know; the pectoral girdle
of a Triassic type recently described by Jackel shows that the
huge acromion, which is one of the most striking peculiarities of
the ordinary form, has arisen in comparatively recent times. There
can be no reasonable doubt but that the girdles of “* Archichelone ”
were of what Prof. Williston calls the “old fashioned” type,
i.e., that there were two coracoidal elements and the pelvis was
“plate-like.” -
Limbs.—From a study of the mode of walking of land Chelonia
it seems to be possible to arrive at some conclusions as to
the type of limbs present in ‘ Archichelone.” In tortoises
the fore arm and leg are carried nearly at right angles to
the upper arm and thigh and stand almost vertical, sometimes
even standing over at the elbow. The humerus and femur are
carried nearly horizontally and very seldom depressed below the
level. The animal sways from side to side, so as to bring its
centre of gravity within the area of support afforded by the three
legs it stands on whilst the fourth is advanced.
Some of these features, the standing over at the elbow for
example, are obviously due to the development of the shell; but,
even if allowance be made for this, there remains sufficient to
suggest that the Chelonia were derived from a group with the
type of humerus found in Lryops and Dimetrodon, which had a
THE CHELONIA. 1015
gait of similar character, the humerus being never depressed
below a horizontal position and moving in a plane which, although
t does fall towards the back, is more nearly horizontal than
vertical. These animals certainly swayed from side to side whilst
walking.
The Chelonian humerus can be directly derived from the
reptilian humerus as represented by Varanosaurus. The rounded
and upturned head is an obvious adaptation to the retraction of
the limbs. The narrow distal end, so different from the wide
epicondyyar region of the early forms, depends on the reduction of
the muscles which lie along the lower surface of the fore arm, and
are inserted on the entepicondyle, following on the habit, induced
by the developing shell, of carrying the fore arm vertically.
The short, powerful, downwardly directed radial and ulnar crests
seem to depend to some extent, though not completely, on the
development of the shell as a source of muscle-attachments.
We may, in fact, take it as probable that ‘ Archichelone” had
limbs more or less like Zryops or Varanosaurus or Dimetrodon.
Hunotosaurus africanus Seeley is at present represented by five
specimens, one of which is in the South African Museum, Cape
Town, and the other four in the British Museum (Natural
History). All these specimens agree in including only the dorsal
region of the animal, and except in the case of the type are
preserved in nodules, so that in transverse section the ribs form
an oval rather higher than wide.
The most nearly complete is No. R. 4054, B.M.N.H. (PI. VII.
figs. 3 & 4). This is preserved in a very hard nodule and shows the
general shape. At the anterior end a scapula is in place on one
side; onthe other end of the specimen we have the impression of
the anterior face of the first sacral ribs and of the anterior border
of the ilia. The specimen is thus of importance, because it shows
that the entire dorsal region is preserved.
No. 49424, which is also preserved uncrushed in a nodule,
is not quite complete behind, but in front the shoulder-girdle is
shown in part. The skull lies with its ventral surface in contact
with the ventral surface of the body and with its anterior end
pointed backwards, with the lower jaw articulated. °
Weathering has. removed the whole skull to within about a
millimetre of the dorsal surface of the palate and in the middle
has even removed the bone from this, leaving a somewhat faint
impression. ‘This specimen is important, because it alone shows
the skull and because the position of the head is only possible if
the neck were fairly long and flexible.
No. 49423 (Pl. VIL. fies, 1 & 2) is a specimen Tote twice the
size of the preceding two, and like them preserved uncrushed.
It shows, well preserved, the whole shoulder-girdle and also im-
pressions of the humeri from which the bone has weathered
away. It also shows traces of scutes.
No. R. 1968. The type-specimen differs from all the otliers in
S. WATSON ON
MR. D. M.
1016
Text-figure 1.
Restoration of Hunotosaurus africanua.
THE CHELONIA,. 1017
that it is preserved in a relatively soft matrix and has been so
macerated before burial that the ribs have twisted round on their
articulations so as to lie nearly flat.
It shows one pubis and a femur, tibia, and fibula,
Skull—The skull is very incompletely and rather badly pre-
served.
The basisphenoid is broad and provided with basipterygoid
processes directed forwards in the plane of its flat lower surface.
There is a rather wide parasphenoid.
The quadrate clearly lies considerably in advance of the occipital
condyle. There are numerous teeth on the palate, some scattered,
but most forming a ridge on the pterygoid. The choane are
rather large and separated by wide bars of the prevomers, lying
close up to the maxille and premaxille.
The maxilla is provided with a single series of small round (?)
teeth, of which about eight are shown.
The premaxille also bear teeth, apparently three in each.
When I first saw the specimen the whole of the extreme
anterior end of the skull was covered by matrix, which I removed
with a needle under a Zeiss binocular dissecting microscope.
Whilst doing so I found no trace whatever of any internarial
processes of the premaxille, and believe them to have been
certainly absent; the anterior nares are consequently confluent
and look directly forward.
The lower jaw is present but not sufficiently well shown for
description.
Except for the evidence given by specimen No. 49424, that the
neck was relatively long and very flexible, nothing is known of
this part.
R. 4054 which, as shown by the position of the limb-girdies,
has a complete dorsal region, has ten dorsals. These are all fairly
similar in structure. The first is short, the second somewhat
longer, and the third very long. The fourth, fifth, and sixth are
about as long as the third, and the seventh to the tenth show a
progressive diminution in length. The structure of the individual
vertebrae is best shown by the type-specimen. The centrum is
very slender, particularly in the fourth to seventh dorsals, and
somewhat hourglass-shaped; it is completely pierced by the
notochordal canal. The rib-facet is carried on a very low and
small process which in the middle dorsal region is placed at the
extreme anterior end of the centrum, whilst anteriorly and
posteriorly it travels back to the middle of its length. The
neural arch is rather massive when compared with the centrum
but is still very narrow. It bears very narrow zygapophyses
which seem to interlock strongly. The upper surface of the
neural arch is essentially flat, the spine being represented only by
a low median ridge. The whole arch appears to be placed very
far forward on the centrum and may overlap on its anterior end.
There is no definite transverse process, but the ribs seem to touch
the sides of the neural arch.
1018 MR. D. M. S. WATSON ON
The first dorsal vib is a narrow, slightly bent, and quite short
bone of an ordinary character. The remaining dorsal ribs, from
the second to the ninth, ave of an extraordinary character. Hach
is strongly curved, articulates by a facet on its proximal end with
the face on the centr um, and then rises, until its upper surface
comes in contact with the neural arch; there is no definite
tuberculum and the capitulum is extremely feeble. The rib
then rises above the level of the neural spine so that the dorsal
surface of the vertebral column lies at the bottom of a groove
formed by the proximal ends of the ribs. The rib now turns
outwards and downwards. The ribs widen very rapidly from
the capitulum, so that until just at the point where they turn
downwards their lateral borders actually touch. They are of a
massive character throughout, and each is strengthened by the
development of a ridge along its visceral surface.
The rib of the fenbe vertebra is not known, but was certainly
not expanded like those which preceded it.
The sacral rib is short and stout.
The shoulder-girdle is well shown in 49423. In this individual
the bones of each side have fused, forming a scapulo-coracoid
which rather strikingly resembles that of Procolophon. The
scapula has a fairly narrow blade with a straight anterior edge,
which has no specialised acromion. ‘The preco‘acoid is a bone
of triangular shape lying flat on the ventral surface. This
results in the scapulze being set not, as usually, in planes parallel
to the principal plane of the body, but inclining together in front,
so that the opening between them towards the neck is very much
narrower than that towards the dorsal region. This implies that
the neck was narrow.
The precoracoid apparently contributes to the glenoid cavity,
and is pierced by a foramen which, exactly as in Pr ocolophon, is
protected by a rounded ridge along g its front border. The
coracoid is a small bone extr emely like that of Procolophon.
The clavicle is a narrow bone running down the entire front
edge of the scapula and then turning slightly in to the inter-
clavicle.
The interclavicle is a long slender rod with a slightly expanded
anterior end which is covered by the inner ends of the clavicles.
The humerus is only incompletely shown.
The nearly complete immature bone in 49424 is badly preserved,
but shows that the bone is narrow and suggests that the deltoid
crest was small.
The impressions in 49423 show that there was a definite head
which is slightly upturned, and that there was an unusually large
but very short ulnar crest.
The pelvis is represented by the hadly preserved impressions
of the anterior borders of the ilia (and pubes 2?) in’R. 4054, and a
perfect pubis in the type. The ilia have a narrow and straight
anterior border and are as high as the space between them. The
THE CHELONIA. 1019
pubis is a small square bone with a foramen, and shows that the
pelvis was plate-like.
The femur of the type is a slender sigmoidally curved bone with
not very well ossified ends and no very marked characters.
The tibia and fibula stand in the specimen at right angles to it ;
in natural articulation thei short stumps present no features of
interest.
Dermal bones are only shown in 49423,
Tn this individual, in the anterior dorsal region a small patch of
what is undoubtedly bone substance is shown lying at a con-
siderable distance above the ribs and neural arches. This is
clearly divided into pieces, one of which forms a small round ridge
in the middle line. The rest of the patch shows a dividing line
running longitudinally and a transverse division apparently
coincident with the line where the two ribs below it meet. On
the posterior end of the specimen, at the same distance above the
ribs, a narrow line of bone is seen in transverse section. This
specimen gives conclusive evidence of the actual presence of
dermal ossifications ; but these are so incompletely preserved as to
make any statements of their distribution of very slight value.
There seem, however, to have been a median series nnd lateral
rows.
The other specimens have no matrix preserved outside the ribs,
so that the scales cannot be seen in them.
The chief features of the structure of Kunotosauria as shown
by these specimens may be summarised as follows :—
The skuli is small, with a wide basisphenoid, a palate of the
ordinary primitive reptilian type with many teeth, teeth in the
maxille and premaxille. The external nares are confluent, there
being no internarial processes of the premaxille. The neck is very
flexible and sharply marked off by its narrowness from the
trunk.
There are ten dorsal vertebre of a slender character with nearly
obsolete neural spines and with rib-facets far forward on the
centra
The first and last dorsal ribs are of an ordinary character, but
the remaining eight are so extremely broadened as to touch one
another by their edges. As shown by the uncompressed speci-
mens, particularly R. 4054, the dorsal region was strongly convex,
and the body was nearly circular in section.
The very small relative size of the pelvis shows that the tail
was separated from the trunk by its much inferior diameter.
The pectoral and pelvic girdles are of ordinary “ old- fashioned ””
type.
The dorsal region is covered with dermal ossifications, of which
there are apparently median and lateral series.
It will be noticed that so far as the structure is known, Huno-
tosaurus agrees exactly with the hypothetical ‘‘ Archichelone”
arrived at by a discussion of the structure of the known Chelonia.
1020 MR. D. M. S. WATSON ON THE CHELONIA.
Not only is this the ease, but in many respects it hasa resemblance
in details even to modern Cheijonia which is very remarkable.
For instance, the long and extremely slender vertebrx are quite
tortoise-like, and in the fact that the capitular facet is placed very
far forward on the centrum we have a suggestion of that inter-
central rib-articulation which is so striking a feature of the
mid-dorsal region of modern Chelonians. The way in which this
rib-articulation travels backwards on the posterior dorsals is
exactly similar to the condition in Homopus. The anterior
position of the neural arch on each dorsal centrum is prophetic of
the modern Chelonian arrangement in which each arch stands on
two centra. The loss of a definite tuberculum on the dorsal ribs
is suggestive of the complete loss of that articulation in Chelonia.
The development of a short, powerful ulnar crest on the
humerus and the slight upturning of its head are also re-
semblances to the same group.
In fact, although our knowledge of Hunotosaurus is too small to
admit of a definite statement to that effect, it is by no means
improbable that it is an actual ancestor of the Chelonia. Whether
it be or not it does, I think, give us a great many suggestions of
the changes which must have taken place during the development
of the Chelonian shell and all that it implies.
T have to thank Drs. A. Smith Woodward and C. W. Andrews
for their kindness to me during my work at the Natural History
Museum, and Dr. L. Péringuey for permission to examine the
South African Museum specimen of Hunotosaurus. The Rev. J.
H. Whaits was the finder of R. 4054, the best existing specimen
of this rare type.
Finally, my discussion of “ Archichelone ” owes much to the
well-known work of many authors, amongst whom Drs. L. Dollo,
O. P. Hay, W. R. Weiland, and J. Versluys are specially
noticeable. When discussing so complex a subject as Chelonian
ancestry it is impossible, if the result is to be at all clear, to
attribute each part to its original author, a fact which must be
my excuse for not acknowledging my indebtedness in detail.
EXPLANATION OF PLATE VII.
Eunotosaurus africanus Seeley.
Fig. 1. Dorsal view of specimen No. 49423. .S., lateral scute; M.S., median
scute.
2. Ventral view of same. Cl., clavicle; Cov., coracoid; H., impression of
humerus ; J.Cl., interclavicle; Sc., scapula.
3. Lateral view of specimen No. R.4054. I7.,ilium: Sec., scapula; I-iX., ribs.
4. Oblique dorsal view of same; I-X., ribs.
All the figures rather less than natural size.
ON SOME CARNIVOROUS THERAPSIDS. 1021
54. Notes on some Carnivorous Therapsids. By D. M.S.
Watson, M.Se., F.Z.S., Lecturer on Vertebrate Pals-
ontology in University College, London.
[Received June 3, 1914: Read November 24, 1914. |}
(Text-figures 1-7.)
INDEX.
Page
Description of Bauria cynops . tek sist on aclawichienionnane ese LOST
x Micr Cooma GURGaCHORE cvadeseueesee LOLS
33 Sesamodon . Jee 1025
Discussion of the Saletoaenine ae the. iBaunides on ine
Cynognathide .............. Smiqiicasnetaen VLOQG
Description of the Brain-case of Seymnognathus sSeaters ewes Ode
e 35 Quadrate region in the Gorgonopsids...... 1034
is 5 Palate of Dycoswehus ..............0--------» 1035
During the last ten years a very large number of perfectly
distinct genera and species of Carnivorous Therapsids from
S. Africa have been described, but of the vast majority of these
we know only the dentition or at most the outside of the skull.
This paper is intended to indicate the very great differences
which may occur in these types, and to show how caution is
necessary in extending any morphological fact discovered in any
form even to its apparently close relatives.
Bavuria cynors Broom, (Text-fig. 1.)
‘Daring my visit to S. Africa I collected from the Cynognathus
zone of Essex, Dist. Albert, a small “ Cynodont” skull; which,
when I found it, was very much weathered and broken into
innumerable pieces. These fit together and give us a nearly com-
plete but somewhat crushed skull, which is curiously preserved,
the bones being very well preserved i in some parts and completely
weathered away in others. The dentition is perfectly preserved.
The skull is considerably smaller than that of the type-specimen
of Bauria cynops, but direct comparison with a cast of that
example and with Dr. Broom’s figures of it, shows no features in
which they differ.
The incisors are large and not of circular section; they are
bluntly pointed, and there is a worn face on the posterior surface
of each, so that they are sub-scalpriform.
The canine is nearly circular in section.
There are ten cheek-teeth visible in the upper jaw, of which
the well-preserved crowns of the first and last four are preserved.
They show no trace of cusps, but are ground down smooth in
such a manner as to suggest a rodent- like movement of the jaw.
There is a short diastema before and behind the canine. :
The lower dentition is not so well known; the lower incisors
are not markedly flattened anteriorly, and do not show clear
1022 MR. D. M. 8. WATSON ON
wear-facettes on their anterior faces corresponding to those which
they undoubtedly made on the upper teeth. They are succeeded
at once by the canines.
The lower cheek-teeth are slightly narrower from side to side
than the upper teeth, and are convex, grinding on the concave
upper crowns.
Text-figure 1.
Bauria cynops Broom.
Restoration of palate, from a specimen collected by the author at Essex,
Dist. Albert, C.P.. x 1.
The palate is quite well shown in the specimen, but in some
regions ouly in impression.
There is a secondary palate, which, however, does not extend
nearly so far back as in Diademodon, and of which the posterior
borders meet in a very acute angle (7. ¢. it has only just closed
up). The posterior nares are divided by a septum, presumably
formed by a median vomer. Posteriorly the roof of the narial
passage slopes down very rapidly. Just behind the last maxillary
tooth on each side the palate is pierced by a large suborbital fossa,
SOME CARNIVOROUS THERAPSIDS. 1023
bounded on the outside by a narrow bar which is certainly trans-
palatine. The pterygoid no doubt combines with this bone to
form the relatively small flange against the inner side of the lower
jaw. Behind this region the two bones are separated by a small
interpterygoid vacuity, which is shown quite conclusively to be
undivided, no parasphenoid crossing it as is usually the case in
early Therapsids.
The inner edge of the pterygoid, which forms the border of
this vacuity, rises, when viewed from below, until it meets its
fellow, and the two then end in suture with the very deep keel of
the basisphenoid.
From the pterygoid at the side of the interpterygoid vacuity
the posterior ramus runs off. It has a deep flange down its outer
border and a horizontal ledge on the inner side. The pterygoid
ramus of the quadrate runs along the front face of this bone as in
all reptiles, whilst in the Cynognathids, in which the apparent
posterior ramus of the pterygoid is really formed by the alisphe-
noid (=epipterygoid), it runs behind.
With the skull of Bauria described above are some very badly
preserved posteranial bones. The scapula is fairly well preserved,
and seems to show that the blade was narrow and had no spine
and no definite acromion.
The coracoid and precoracoid are also shown; they are of the
ordinary South-African Therapsid type, but are relatively very
large. The whole arrangement must have closely resembled that
figured by Dr. Broom in /etidosaurus.
MiICROGOMPHODON OLIGocyNus Seeley. (Text-fig. 2.)
Another Cynodont which must be regarded as a close ally of
Bauria is Microgomphodon oligocynus, known only by the type-
skull. [The fine skeletal fragment which Seeley referred to this
-genus is shown by the very unsatisfactory jaw fragment to he
quite distinct, being, no doubt, a Cynognathid. |
ape type-skull is illustrated with slight restoration in text-
fig. which should be compared with Prof. Seeley’s excellent
Tcheewenie figures (Phil. Trans. 1895, B, pl. i. figs. 1-4). The
animal resembles Bawria in the following respects :—
The short temporal region.
The relatively heavy face.
The dentition.
The presence of an interpterygoid vacuity.
The presence of suborbital vacuities.
The slight reduction of the posterior part of the lower
jaw.
7. Small squamosal,
8. The nostril directed more anteriorly than outward.
9. The nasal not widened. posteriorly.
10. The frontal entering into the orbital margin.
SOU oe bo
The upper incisors are subscalpriform and worn, exactly as in
1024 MR. D. M. 8. WATSON ON
Bauria, and the very large and procumbent lower incisors are also
very markedly chisel-shaped. There seems to be little doubt but
that the wear of these teeth is similar to that of rodent incisors
and that the front of the tooth was harder than the back, although
the teeth seem to be enamel-covered all round.
Text-figure 2.
Microgomphodon oligocynus. X 1.
A. Restoration of skull from the dorsal aspect. From Prof. Seeley’s
ty pe-specimen.
B. Restoration of skull from the side.
A curious feature of this type, shared also but in a less marked
degree by Bauria, is the very great space between the posterior
ramus of the pterygoid and the paroccipital process.
SOME CARNIVOROUS THERAPSIDS. 1025
SESAMODON.
Another type which has to be brought in connection with
Bauria is Sesamodon. This type is probably represented in the
British Museum by the anterior part of a skull broken off through
the middle of the orbits. Part of the dentition is fairly well
preserved. The skull, so far as it goes, is of exactly the same size
and proportions as the type, but differs markedly from Dr. Broom’s
restored figure in that the lower canine does not bite outside
the maxilla but inside, in the usual way among Therapsids. As
Dr. Broom’s type-specimen is extremely weathered and badly
preserved, it is quite possible that the part of the maxilla outside
this tooth was removed by weathering.
In any case, judging only from Dr. Broom’s figures of the type-
specimen, the animal resembled Lawria in the following ways :—
Short temporal region.
The heavy face.
The general features of the dentition.
The small squamosal.
The feeble postorbital arch.
The frontal forming part of the orbital margin.
The nasal not widened posteriorly.
The prefrontal excluding the lachrymal from the nasal.
The interpterygoid vacuity, and the great distance between
the posterior ramus of the pterygoid and the paroccipital
process.
COON A Tp oo bo
The group, which may be called the Bauridee, including these
three reptiles, differs from the Cynognathide (Cynognathus, Dia-
demodon, and Trirachodon) and the Nythosaurid in the following
series of characters :—
1. The short temporal region.
[2. The heavy face. ]
3. The nostril directed more forward than outward.
The nasal not widened posteriorly.
The frontal forming part of the orbital margin.
The powerful incisors and grinding “ molars.”
. The small squamosal.
8. The large septomaxille on the face.
9, The interpterygoid vacuity.
10. The suborbital vacuity.
11. The great distance between the posterior ramus of the
pterygoid and the paroccipital process.
lla. The absence of the long narrow bar formed by the
pterygoids and parasphenoid behind the region of the
transpalatine.
12. The presence of a quadrate ramus of the pterygoid.
13. The posterior position of the notch in the angular and the
large size of its reflected lamina.
14. The lack of an acromion and spine on the scapula.
15. The large coracoidal elements.
Proc. Zoou. Soc.—1914, No. LXIX. 69
(SU)
1026 MR. D. M. S. WATSON ON
"hese characters are, of course, of very different values, but,
taken together, do show that the Bauridee differ enormously from
the Cynognathide.
It may be argued with an appearance of truth that these
characters are all merely primitive ones, and that the Bauridee
are derived from the same near ‘“ Cynodont” ancestor as the
Cynognathids, but, although much specialised in some ways, are
retarded in other features. This is in essence the view that
Dr. Broom stated of Bauria in the form of a genealogical tree
in December 1911, and which, I believe, he still holds.
IT endeavour to show in the remainder of this paper that
Dr. Broom’s view is now untenable. ‘The method I employ
is to discuss, with the aid of a series of specimens, of which the
relative ages ave known, the evolution of certain selected features
of the skull of a Gynognathid, and then endeavour to show that
the conditions in the Bauridee could not have been derived from
those in any, except perhaps the earliest, of these stages.
The series of forms | shall use are :-—
Dimetrodon; Upper Carboniferous ¢ or Avtinskian.
* Arctops willistoni, gen, et sp. n.; Hndothiodon zone? Middle
Permian ¢ or earlier.
Scymnognathus whaitsi? Broom; Hndothiodon zone. Middle
Permian.
Arctognathus curvimola (Owen); Cisticephalus zone. Upper
Permian.
Diademodon; Cynognathus zone. Middle Trias.
The first of these is a North-American Pelycosaur, and the last
a South-African Cynognathid. The other three are South-African
Gorgonopsids. It is certain that they are not actual ancestors of
one another, but they may be as nearly related as the animals
included in the early phylogenies of the horse.
Baur, Case, and v. Huene have published more or less complete
accounts of the occiput of Dimetrodon, and I am indebted to the
extreme kindness of Prof. 8. W. Williston for the gift of a magni-
ficent specimen which has been of the utmost use to me. Dinetro-
don has a large, round occipital condyle, in front of which the
thick basioccipital forms the narrow floor of the brain-cavity, and
articulates at the sides with the exoccipitals and opisthoties,
which are usually distinct. The opisthotics form powerful par-
occipital processes lying below the small post-temporal fossz.
The small exoccipitals lie on the extreme posterior surface,
projecting at the sides into processes which run outwards towards
the post-temporal fosse far above the bottom edge of the par-
occipital processes. The whole bone looks as if it had only
recently become a part of the skull, and was still only very
imperfectly connected with the rest of the occiput. The foramen
jugulare lies high up at the back of the skull between the lower
* Wounded ona skull in theG. G. Bain collection, B.M.N.H. R. 4099, from Howse’s
Port. It is distinguished by the extreme width of the parietal region and the small
size of the orbit.
SOME CARNIVOROUS THERAPSIDS. 1027
edge of the lateral process of the exoccipital, the opisthotic, and
the basiocecipital.
The front face of the paroccipital process is formed by the
pro-otic, and its lower surface is channelled. ‘The fenestra ovalis
is a large, irregular hole, not bounded by bone in front, lying
below the level of the base of the basioccipital condyle. The
special process of the opisthotic which forms its posterior border
has a wide, smooth face looking forwards and outwards, and is
supported on the inner side by a corresponding process from the
basioccipital.
The basisphenoid, as shown by Case’s figures, has large tuber:
reaching back towards the fenestra ovalis, and powerful basi-
pterygoid processes of a more or less ordinary character.
The basicranial and otic region of the very primitive Gorgon-
opsid Arctops closely resembles that of Dimetrodon. It has a
large, round basioccipital condyle and a thick basioceipital which
supports the exoccipitals and opisthotics. The paroccipital pro-
cesses are very powerful, lying below the small post-temporal fosse.
The exoccipitals are extremely similar to those of Dimetrodon,
lying on each side of the foramen magnum and sending a special
process out towards the post-temporal fossee. The exoccipital
even in this type is only incompletely included in the occiput.
The foramen jugulare is high up on the back of the skull.
The front face of the paroccipital process is formed by the
pro-otic, and its lower border and front face are channelled by
a groove at the inner end of which lies the large and rather
irregular fenestra ovalis. This is placed at about the level of the
lower surface of the basioccipital condyle, and is completely
enclosed by bone.
The basisphenoid forms two very large tubera, between which
is a deep depression terminated abruptly in front by the junction
of ridges from the tubera to form a very deep median keel con-
tinuous with the parasphenoid.
The tubera basisphenoidalia are far back, and end in a ridge
just in front of the fenestra ovalis.
The basipterygoid processes are curious horizontal plates
standing out from the sides of the flat, vertical plate which
forms the whole anterior part of the basis cranii.
Scymnognathus whaitsi* differs far more from Aretops than
the latter does from Dimetrodon in this region.
The basioccipital condyle is no longer round, but forms a mere
lip round the lower border of the foramen magnum. The whole
arrangement is so remarkable that if I had not myself removed
the thin skin of matrix surrounding it, I should not have believed
it possible. This lip is no doubt partly formed by the exoccipital,
as it extends up round the sides of the foramen magnum.
* This is the specimen the lower jaw of which I described under the name
Seymnosuchus whaitsi, Ann. & Mag. Nat. Hist. ser. 8, vol. x. p. 578, fig. 3. Iam
not certain of the identification.
69*
1028 MR. D. M. 8S. WATSON ON
The basioccipital is a very thin bone, articulating as usual with
the exoccipitals and opisthotics.
The exoccipital is an exactly similar bone to that of Arctops.
The paroccipital process is very shallow, so that the foramen
jugulare lies much nearer the bottom of the skull, and actually
looks as much downward as backward. The front face of the
paroccipital process is slightly channelled, and the fenestra ovalis
lies at the inner end of this groove above the level of the basi-
occipital condyle. It is a comparatively small hole surrounded
by a smooth and badly-preserved area of bone.
The tubera are much lower and much further forward than in
Arctops, the lower surface of the basisphenoid being only com-
paratively slightly depressed.
The basipterygoid process is very similar to that of Arctops,
but seems to be pierced by a canalis Vidii.
It is at once obvious that most of the differences between this
type and Arctops are due to a reduction in depth of everything
which lies below the bottom of the foramen magnum.
The type and only existing skull of Arctognathus is unfortu-
nately damaged so that nothing behind the fenestra ovalis is
visible. In this form the basioccipital is even thinner than in
Seymnognathus, and the fenestra ovalis lies above the bottom of
the foramen magnum. The tubera basisphenoidalia are repre-
sented only by the thickened margins of the slightly concave
triangular lower surface. The basipterygoid process is similar to
that in Seymnognathus, but is relatively further forward.
In Diademodon the basioccipital is so thinned that the centre
part of the thin lip-like condyle of Seymnognathus is pinched out
altogether, leaving the two condyles, which are no doubt mainly
exoccipital. Although no definite suture has ever been seen, the
texture of the bone makes it certain that the exoccipital in
Diademodon is a small triangular bone forming the lower margin
on the extreme back of the skull. The fenestra jugularis les
entirely on the lower surface and does not face in the least
backward. The paroccipital process is more massive than in
Scymnognathus owing to the fact that the post-temporal fosse lie
above the level of the foramen magnum. ‘The front face is
channelled, and the fenestra ovalis les at the inner end of the
groove considerably above the lower margin of the foramen
magnum.
The tubera basisphenoidalia are very reduced, being merely
the thick edges of the nearly flat ventral surtace of the triangular
basisphenoid.
The horizontal basipterygoid process is similar to that of
Scymnognathus, but much further forward.
Arctognathus is in all visible features of this region exactly
intermediate between Scymnognathus and Diademodon.
Comparison of these descriptions with text-figs. 3 and 4 will
show that the changes take place quite regularly with time, and
that all of them depend eventually on the reduction of the
whole regions which lie below the base of the brain, a reduction
SOME CARNIVOROUS THERAPSIDS. 1029
which is shown extremely clearly in the three drawings of text-
fig. 3.
Text-figure 3.
Occipital views of the skulls of :—
A. Diademodon. |
C. Arctops willistoni, gen. et sp. n.
Foramen jugulare ;
Interparietal ; B.Sp., Basisphenoid.
B. Scymnognathus whaitst ?
Par.Oc., Paroccipital process ; XS: Fixr.Oc., Exoccipital ;
Sq., Squamosal ; Tab., Tabulare; I.Par.,
1030 MR. D. M. 8S. WATSON ON
Now, according to Dr. Broom’s figure of the occiput of the
type-skull of Bauwria, the basioccipital condyle is large and
rounded, much more like that of Avctops than any other of the
series described above. The fenestra lies far above the bottom
of the occiput, looking directly backwards. There is no trace of
the gradually developed triangular lower surface of the basi-
sphenoid which is so characteristic a feature in Cynognathids.
In fact, it seems quite certain that the basicranial region of
Bauria is the product of an utterly different type of change from
that which has led to that of Diademodon, and as the features of
Text-figure 4.
WW
i
HI
q
Await PEN 2 (i
PNM
Auli tl(l
NG
The occipital and otic regions of :—
A. Dimetrodon.
B. Arctops willistoni.
C. Seymnognathus whaitsi ?
D. Diademodon.
Viewed from below and the right side.
Tub., Tuberosities of basisphenoid; Oc.C., Occipital condyle; X., Foramenjugulare ;
B.Pt,, Basipterygoid process ; 7.Ov., Fenestra ovalis,
SOME CARNIVOROUS THERAPSIDS. 1031
this region in the Hndothiodon-zone Gorgonopsid Scymnognathus
are obviously prophetic of those of Cynognathids, it will follow
that the Bauride have had a quite different ancestry since that
time, which is long anterior to the development of a ‘‘Cynodont ”
structure—7. e, a secondary palate in any form.
This conclusion, if true, will necessitate the splitting up of the
Order ‘* Cynodontia,” and the recognition that it includes at least
two distinct branches which were separate when they indepen-
dently acquired a secondary palate.
The detailed structure of the palate of any Gorgonopsid is
not known beyond the possibility of doubt.
All Gorgonopsids which I have seen agree in the following
features :—
lst. There is a long region in front of the basisphenoid in
which the two pterygoids are in contact, apparently with a
median parasphenoid, but perhaps for part of the distance with
each other, and in which their lateral margins are parallel ;
that the whole forms a narrow bar behind the powerful descending
processes of the pterygoids.
2nd. If an interpterygoid vacuity is present it is very small,
and lies just between the descending flanges of the pterygoids.
3rd. There is a deep groove down the middle of the posterior
part of the palate.
Ath. There are no suborbital vacuities.
5th. There are very large posterior nares, which are separated
by a bar lying considerably above the level of the lower edges of
the maxille, which tend to be approximated.
No skull I have seen shows all the sutures on the palate satis-
factorily. The skull of Arctops shows many of them, but most
unfortunately, owing to its having been split longitudinally,
those in the middle are not clearly visible posteriorly.
The type-skull of Goryonops shows many sutures in a rather
indefinite manner.
As Dr. Broom has pointed out. the bar which divides the pos-
terior nares in Gorgonops and many othe: forms is actually single.
It has to be considered whether, like the single *‘ vomex” of the
Anomodonts, it 1s really composed of a pair of fused prevomers,
or whether it is a parasphenoid. Its complete resemblance in
shape, even to the grooves on its upper surface for Jacobson’s
organs, to the undoubted prevomers of many Therocephalia,
suggests that it is to be interpreted in the same way; and in the
type-skull of Arctops it actually seems to be double when seen in
section on the anterior end of the specimen, about one centimetre
in front of the posterior end of the posterior nares. In this
specimen, in place of the single dorsal ridge which occurs in
Scymnognathus, there are two recs than a millimetre apart, and
the groove between them seems to be continued by a suture into
the palate. If this observation is correct, and the condition of
the specimen in this region is not good enough for certainty, then
there can be no doubt Siipats the bar between the palato-nares in
1032 MR. D. M. S. WATSON ON
Gorgonops is really composed of a pair of prevomers fused, just as
is the rather similar bone in the Anomodonts.
In the type-specimen of Gorgonops it seems clear that there is
no median suture down the palate between the posterior nares and
the very small interpterygoid vacuity. On the other hand, there
seems to be very indefinite sutures along the side walls of the
median groove of the palate, exactly in the position where the
sutures between the vomer and palatines and pterygoids lie in
Diademodon.
Tf this: be so then Gorgonops will have a large mammalian
vomer in the posterior part of its palate and a fused pair of pre-
vomers anteriorly.
In Aretops, which is much more primitive than Gorgonops,
there is some evidence suggesting that the pterygoids reached
forward to the prevomers.
Comparison of the series of characters recorded above as
features of the Gorgonopsid palate with the similar series on
p- 1022 of this paper referring to Lauria, will show that Lauria
differs in its palate in exactly the same manner from Gorgonops
as it does from Diademodon.
In fact the Gorgonopsid palate, as I have previously pointed
out, presents a very close resemblance to that of the Cynognathids,
a resemblance which is much more striking in the advanced
Arctognathus than in the more primitive Gorgonops.
If the interpretation of the structure of the palate of Gorgonops
given above be correct, then the only differences between it and
that of a Cynognathid are the development of a secondary palate,
the beginnings of which are seen in that type, and the concurrent
reduction of the prevomers; a change which is paralleled in
Crocodiles.
Amongst the other characters, of which a progressive and
orthogenetic change is seen in this series of skulls, are the occiput
and the temporal region.
The occiput of Arctops (text-fig. 3C) is very nearly flat, with
a minute foramen magnum and only very slight projecting ridges
formed by the tabulares and squamosals. It only differs from
that of the Deinocephalian Z%itanosuchus in the slightly greater
spread of the squamosals. In particular, it resembles this animal
in the very great breadth of the interparietal.
In Scymnognathus (text-fig. 3 B) the occiput is deeply concave
and the squamosals very wide. The interparietal is very much
narrower than in Arctops.
In Diademodon (text-fig. 3 A) the occiput is very concave and the
interparietal narrow. This form, however, differs pronouncedly
from the Gorgonopsids in its very largely developed squamosals,
a feature not shown in the Nythosaurians which seem to be
related to it.
In the temporal region Arctops is remarkable amongst Gorgon-
opsids for the extraordinary width of the parietal region and the
[s))
SOME CARNIVOROUS THERAPSIDS. 103
shortness of the temporal fosse, which are only about half as long
as the width of the parietal bone.
In Scymnognathus the parietal region is much narrower, and
the temporal fossz nearly as long as the parietal region between
them is wide.
In Arctognathus the temporal fosse are longer than the parietal
region 1s wide, and in the Cynognathids they are very long, and
the parietals form a narrow crest.
Brain-cavity of Scymnognathus.
The specimen of ‘ Seymnognathus” shows the brain-cavity,
although, owing to the hardness of the matrix with which it was
filled, this is not very well preserved. Its main features are,
however, quite certain, but it does not merita lengthy description.
The general structure will be best understood from text-fig. 5.
Text-figure 5.
The brain-cavity of Scymnognathus whaitsi ? in sagittal section. X 1.
There is a distinct resemblance to Diademodon owing to the
comparatively thin basioccipital and the large opening to the
vestibule through the posterior end of which the tenth nerve had
its exit. The general features of the vestibule recall Diademodon,
but there is no visible trace of a cochlea.
The chief differences from the more recent animal are that the
whole cavity for the cerebellum is very much smaller, and that
its bage rises very rapidly in front, very much as it does in
Dimetrodon.
The prootic, although it extends further forward than in the
Anomodonts and Dimetrodon, has more of the greater anterior
1034 MR. D. M. S. WATSON ON
projection which, oceurring in Diademodon and also in Ornitho-
rhynchus, has a long suture with the parietal.
Quadrate region of Gorgonopsids.
A large Gorgonopsid skull, at a slightly more advanced stage of
evolution than Arctops, from the Hndothiodon zone of Beaufort
West, which was collected by the Rev. J. H. Whaits and is now
in the British Museum, shows the quadrate and the bones
surrounding it in a most perfect manner.
Text-figure 6.
The quadrate and surrounding bones of a Gorgonopsid from the
Mndothiodon zone of Beaufort West. X 1.
A, from in front; B, from behind.
Par.Oc., Paroccipital process; Pt., Pterygoid; Quw., Quadrate; Qu.J., Quadrato-
jugal ; Sqg., Squamosal; Tab., Tabulare.
The squamosal has a powerful articulation with the end of the
paroccipital process ; the bone there sends a process dorsally and
medially which articulates with the tabulare, postorbital, and
SOME CARNIVOROUS THERAPSIDS. 1035
probably the supraoccipital and parietal. Lateral to its articu-
lation with the paroccipital process the squamosal is very massive,
and after passing directly outward for some distance finally
curves round and forms part of the zygomatic arch. The front
face of the lateral part of the squamosal is excavated for the
quadrate and quadratojugal.
The quadrate is a comparatively large bone with a trace of the
pterygoid ramus in a slightly turned back area behind which the
posterior end of the pterygoid passes. The lower margin of
the bone is thickened to form the condyle, which, although its
surface is destroyed, must have looked very much more forward
than downward.
The quadratojugal is a small bone completely fused with the
quadrate at the lower margin and actually forming a good deal
of the condyle. On the posterior surface it is seen to overlap the
quadrate ; separating a good deal of its posterior surface from
the squamosal between the two bones is a large foramen.
In Arctognathus the quadrate is partially shown on one side,
and relatively is considerably smaller than that of the earlier
type described above.
In Diademodon the quadrate as it has been described by
Seeley, Broom, and myself, is very much smaller. It will be
remembered that 1t is possible that there is a foramen through
the quadrate of Diadenodon suggesting that even in that form
there is a rudimentary and otherwise totally fused quadratojugal.
The Palate of Lycosuchus ?
In the British Museum there isa very large “ Therocephalian ”
skull (R. 4100) from Vit. Kyk. Gouph, collected by T. Bain from
the Tapinocephalus zone.
This is perhaps not definitely determinable, as it is broken
off through the canine, behind which it shows only one molar
tooth. I can see no generic differences, in fact few differences,
except that it is considerably larger, from the specimen of
Lycosuchus vanderriti described by Dr. Broom. In any case it
is unquestionable that it is closely allied to Scylacosaurus. The
palate of this individual is very well but curiously preserved,
having been cleaned by the weathering of a broken face which
passes along the palate, having its mediai portion adherent to one
block and the remainder to another.
The basisphenoid is not completely shown, but has a very deep
(2 ecm.) and narrow keel along its ventral surface. Although
they are only incompletely exposed, iteis evident that the basi-
pterygoid processes are the usual flat plates found in all South
African “Carnivorous” Therapsids. It is clearly shown that the
epipterygoid, which is a comparatively narrow but laterally
flattened bone, articulates with the process.
The pterygoid is the usual large triradiate bone ; it articulates
with the basipterygoid process and sends a ramus pack to pass
1036 MR. D. M. S. WATSON ON
behind the quadrate. The ramus has a ridge down its outer
margin and a flat shelf on the inner side.
Text-figure 7.
Restoration of the palate of Lycosuchus? X oe
B.Sp., Basisphenoid ; P.V., Prevomer ; Pal., Palatine; Pt., Pterygoid ;
Vo., Vomer.
In front of the basisphenoid the pterygoid rises into a high
ridge overlapping on to the basisphenoidal keel, which terminates
SOME CARNIVOROUS THERAPSIDS. 1037
suddenly, leaving the ridge on the pterygoid to form the side of a
narrow interpterygoid vacuity ; further forward the flange of the
pterygoid meets its fellow of the opposite side, and the two run
on in contact till they are again separated by a small round
interpterygoid vacuity.
At the side of this region the process which combines with
the transverse bone to form the powerful pterygoid flange is
given off.
Further forward the bone bears a ridge with two irregular
rows of teeth, and finally undoubtedly continues forward to the
prevomer.
On the block from which the median part of the palate has
been split away, there is very clearly shown a long. median bone
forming a narrow vertical plate. That this bone is not, as it
might conceivably have been, formed by a fused pair of ridges on
the dorsal surface of the pterygoids, is shown by the occurrence
of very fine but I think definite sutures, between its posterior
end and the pterygoids, and conclusively by the fact that what is
undoubtedly a part of the pterygoid is applied to its lateral face
ending in front ina very obvious manner. ‘This median bone is
clearly seen to pass between the posterior ends of the prevomers
in front.
There can be no doubt but that this bone is the same as the
vomer of Diademodon, and is the median bone in the same position
which, if Lam right, occurs in Gorgonops.
The palatine is a large bone with a long suture with the
maxilla; posteriorly it forms the front border of the large sub-
orbital fossa and its inner edge has a suture with the pterygoid,
finally it has a short contact with the prevomer.
The two prevomers are very clearly seen in the specimen ;
posteriorly they are separated by the vomer, but in front they
are in contact ; the lower surface of the bar between the posterior
nares, which is formed by them, has exactly the same form as that
of the similar bar (which in my view is probably formed by a
pair of fused prevomers) in Glorgonops.
The palate just described differs from that of Scylacosaurus
only in the presence of the very narrow vomer.
This palate will be seen to resemble that of Lauria in very
many features; in fact it differs from the palate of the nearly
contemporary Gorgonopsids in exactly the same features that the
palate of Bauria differs from that of a Cygnognathid.
This specimen is the first that has shown a definite median
vomer and a pair of paired prevomers in the same animal. It
seems to me to add a very strong argument in favour of Dr.
Broowm’s view that the mammalian vomer is not homologous with
the “vomers ” of a Lizard.
The various new facts brought forward in this paper show that
the accepted division of the “ Carnivorous” Therapsids into
Therocephalia, Gorgonopsia, and Cynodontia, is far from satis-
factory. In my opinion the differences between Lauria and its
1038 ON SOME CARNIVOROUS THERAPSIDS,
allies and the Cynognathide are important—are, in fact, to a
great degree actually the same as those separating the “ Thero-
cephalia ” and “ Gorgonopsia.” I do not regard these differences
as being of ordinal value, and as a purely temporary measure
propose to revive Owen’s term of Theriodontia as an Order,
including in it as suborders the Therocephalia, with Scylagosaurus
and Lycosaurus as well-known types; the Gorgonopsia, with
Gorgonops, Arctops, Scymnognathus, Arctognathus ; the Bauride,
with Bauria, Microgomphodon, and Sesamodon; and the Cyno-
dontia, to include the Nythosauride: and Cynognathide.
ON A NEW MAMMALIAN GESTODE. 1039
| From the Procnupines or rue Zoontoaican Socinry or Lonvon,
1914. ]
{Published December 1914.
Contributions to the Anatomy and Systematic Arrange-
yy y gs
ment of the Cestoidea. By Frank EH. Bepparp, M.A.,
D.Sce., F.R.S., F.Z.8., Prosector to the Society.
(Text-figures 1-9.)
XV. On A NEW GENUS AND SPECIES OF THE
Famity AcoLEIDa.
INDEX. Page
Moneecocestus erethizontis, gen. et sp. 0. ........0.--00000--.2--. 1055
T obtained a large number of examples of this new Tapeworm
from a specimen of the Canadian ree-Porcupine, Hrethizon
dorsatum, which died in the Society’s Gardens on May 14. From
the comparatively short time which the host had lived in the
Gardens (one year and two months) it would seem to be very
possible that it was infected with the parasite before arriving in
London, and that this Cestode is therefore an American and not
a Kuropean genus or species. On the other hand, there are no
facts which would render the opposite view untenable. But little
is known of the longevity of these worms in their final hosts.
To the naked eye, or after an inspection with a hand-lens,
there is nothing remarkable in the worm, which, however,
shows on a more profound anatomical examination to present
several features of interest, of which the most important is the
great reduction of the female efferent organs. This feature
would seem to place it inthe family Acoleidz ; but the systematic
position will be considered after the structure has been gone
into.
This species is medium sized, the examples reaching a length
of 50 to 60 mm., or perhaps a trifle more; the greatest breadth
is not more than just over 3 mm. When alive, the scolex end
of the worm swayed about with considerable vivacity, which
I have not observed to be generally the case among tapeworms,
the movements being, as a rule, slower.
The segments are flat and thin, and at the end of the body get
to be rather translucent. This lack of the usual white appearance
is not, however, due to sterility ; the very last segment in such
examples as I examined, by means of sections, was gravid. The
scolex is very distinct, though the body widens immediately ,
after it. There are no hooks of any kind upon either the
rostellum or the suckers. There is, im fact, no rostellum; the
suckers are fairly large and conspicuous. Under examination
with a lens, the neck appeared to be quite absent and the seg-
mentation to commence at once. There is, however, as may be
seen 1n longitudinal sections, a very short neck where no segments
1040 DR. F. E. BEDDARD ON
are to be observed. In the strobila region the segments ‘are
at first very narrow; but they increase in length posteriorly,
though never reaching a greater length than diameter. The
hinder margins on either side project in the more mature
segments over the following segment. The last segment of the
body was often to be seen; it was narrower and a trifle longer
than those immediately preceding it and somewhat oval in form.
As already remarked, this segment did not prove to be sterile.
Text-figure 1.
LeSs
Moncecocestus erethizontis.
A. General view of scolex and anterior proglottids.
B. A few anterior proglottids, illustrating relative positions of receptaculum
seminis and cirrus-sac.
p. Genital pore. 7.s. Receptaculum seminis.
The genital pores alternate with rigid regularity from side to side
of the body. The cirrus was frequently seen to be protruded,
and very often an oval body bearing the cirrus also projected ~
trom the side; on making sections this body proved not to be the
protruded cirrus-sac as might have been anticipated, but that
sac surrounded by a certain part of the medullary tissue which
together formed a marked bulge upon the side of the segment.
A NEW MAMMALIAN CESTODE. 1041
These are the only external characters to be noticed in this
worm, whose identity or systematic position, as is obvious, could
not be thereby determined. The flatness and thinness of the
strobila is emphasised by transverse sections. In such, the fact
may be observed that the cortex is of less diameter than the
medulla. As is usual, the two layers of parenchyma are separated
by a layer of transverse muscles, which is, in the present species,
Text-figure 2.
Monecocestus erethizontis.
Part of a transverse section through a ripe proglottid.
ci. Cirrus-sac. ov. Ovary. 7.s. Receptaculum seminis. w¢. Uterus.
w.v. Water-vascular vessels.
a thin one. So, too, are the longitudinal muscles, which are no-
-where massed into bundles. They lie singly, but in several rows ;
nor is their arrangement regular. Thus the muscular system is
altogether slight, and contrasts in this with various members of
the family Acoleide. The muscular fibres themselves are slender,
Proc. Zoou. Soc.—1914, No. LXX. 70
1042 > DR. F. E. BEDDARD ON
and it is not a little unusual to find that: the abundant dorso-
ventral fibres are of stouter build, and thus much more
conspicuous in a transverse section. Calcareous bodies are
numerous in this species, and are to be found throughout the
body both in the cortex and the medulla.
The water-vascular system consists of the usual four longi-
tudinal trunks, of which the ventral are connected by the usual
transverse vessels in each segment. Both in transverse and
horizontal sections it can be observed that the two trunks of each
side of the body are placed as nearly as possible side by side.
The outermost of the two vessels, however, is slightly to one side,
that is to say, it is, being the dorsal vessel, slightly dorsal in
position of the ventral tube. It is well known that the relative
positions of the dorsal and ventral tubes when lying side by side
differ, being in some species arranged precisely in the reverse
fashion to others. In the present worm the dorsal vessel lies to
the outside, as I presume from the fact that the outer vessel is
not the one which is connected with its fellow by the transverse
vessel in each segment, and from the fact that it is from time to
time much narrower in calibre than the inner ventral vessel.
Often, however, the two vessels are of the same size. The trans-
verse tubes lie in the middle of the medulla, and at the end of
the segment. The inner longitudinal vessel bends inwards to
give off the transverse vessel, which thus forms a Y-like angle.
I could discover no trace of any network connected with any of
the main trunks. Such a network exists in the family Acoleidee,
and is referred to by Fuhrmann * in Shipleya inermis.
nd
Genital Organs.
A peculiarity of this species—which, however, it shares with
a good many Cestodes—is the early development of the genital
organs. It agrees, for instance, with Schizotenia americana (to
accept v. Janicki’s identification of Stiles’s species tT) in this
feature, a point which must be borne in mind in comparing the
two. I recognised in longitudinal horizontal sections consider-
able traces of the genitalia in the fourth segment, and I would
not like to assert the absence of still more undifferentiated
beginnings in an earlier segment. In the sixth segment the
cirrus-sac was fairly developed and the sperm-duct obvious ; and
in the sixth segment I saw for the first time a perfectly clear
circular external pore. This was seen in a specimen mounted
entire in glycerine, and it agrees with another specimen which
I examined by sections. There were pores in all the segments
posterior to the sixth. I have already mentioned the absolute
regularity of the alternation of the genital pores from right to
left side. The earliest segment in which the genitalia appeared
to be quite functional was the ninth, in which J observed the
* Rey. Suisse Zool. xvi. 1908, p. 70.
+ Proc. U.S. Nat. Mus. xix. 1897, p. 165.
A NEW MAMMALIAN CESTODE. 1043
receptaculum to be full of sperm. Neither the testes nor the ova
were ripe so early in the body.
The testes lie posteriorly in the segment just in front of the
transverse water-vascular vessel. They form a narrow band
running right across each segment, but contained within the
area bounded by the lateral water-vascular trunks.
It may be seen in transverse and horizontal sections that the
testes are not more than two deep and are in three or four
rows. There are about 60-70 in each segment. I regard the
testes as dorsal in position, because when the outer and dorsal
water-vascular trunk moves away from its lateral position with
reference to the inner tube, it les to the side corresponding to
that on which lie the testes. They are therefore dorsal, and the
ovary 1S ventral in position.
Furthermore, since the cirrus-sac passes to the same side of the
two water-vascular tubes as that on which the testes lie, it has a
dorsal position with regard to them ; this point is not mentioned
by Fuhrmann in his account of Shipleya. Moreover, as the trans-
verse vessels arise from the ventral vessels they, too, are ventral
in position, and therefore the female organs such as the recep-
taculum, which is more or less level with them, must be ventral.
Fuhrmann states that in Shipleya the ovary is ventral. JI have
not been able to notice any ripe masses of spermatozoa in the
testes, though there are many in the cirrus. This absence of
sperm from the testes has already been noted by Fuhrmann in
another genus.
The vas deferens of this worm is very remarkable. In the
very young proglottids the vas deferens emerges from the cirrus-
sac asa short and curved tube which is curved backwards towards
the testes. In rather older proglottids the vas deferens is not
much longer but is distinctly differentiated into two regions;
there is a ‘wider tube which emer ges from the cirrus-sac and this
abruptly becomes narrower in the distal region, where it ulti-
mately breaks up into three or four branches for the supply of
the testes. This differentiation of the vas deferens is much more
highly developed in mature proglottids. The vas deferens after it
emerges from the cirrus-sac runs forwards and inwards obliquely
towards the anterior end of the segment, and that is away from
the testes, which are posterior in position ; this region of the vas
deferens is that of the greater calibre, and the fine tube which
issues from it is the sperm-duct proper and runs backwards
towards the testes and thus at an acute angle with the anterior
region. This is foreshadowed in the curved course of the entire
vas deferens shown in the immature proglottids. To return to
the anterior section of the vas deferens—this tube is very wide,
relatively speaking, and often dilates at the ends, where it passes
into the narrow section of the vas deferens, into a circular sac ;
its width varies in parts and according to the state of develop-
ment of the segment.
The intimate structure of this region of the vas deferens is a
(OTs
1044 DR. F. E. BEDDARD ON
little difficult to make out. But it is clear that it forms a tube,
slightly sinuous, which is walled by a layer of cells in which
appear to lie masses of secretion, which masses are colloid in
appearance and deeply stained by hematoxylin. They are found
also within the tube, especially in its more dilated regions.
These masses of secretion, as I suppose them to be, are shown
Text-figure 3.
Monccocestus erethizontis.
A. Cirrus-sac in longitudinal section.
a. Its cellular sheath. ce. Slightly produced cirrus.
B. Proximal end of vas deferens.
ci. Cirrus-sac. 7. Longitudinal muscular fibres. v.s. Dilatation of
vas deferens.
in text-fig. 3. The tubular character of this dilated region of
the vas deferens is very obvious in the younger proglottids, where
the cells of the tube have not become modified by the secretion
spoken of. In parts, for instance, in the terminal dilatation
A NEW MAMMALIAN CESLODE. 1045
represented in text-fig. 3, the walls of the tube are thin but
apparently still cellular. From this dilated extremity springs
the fine and narrow distal region of the vas deferens, in which
the actual walls are to be recognised with difficulty. These
appearances suggest that we have to do here with an extremely
exaggerated form of a vesicula seminalis, which possibly serves
other purposes beside the mere storage of sperm; but as to what
these functions are I have no reasons to form an opinion. It
is, however, perhaps to be associated with the development of
the sperm, which does not appear to come to maturity within
the testes, as I have already pointed out. In all proglottids the
dilated and glandular region of the vas deferens appeared to be
full of sperm.
The cirrws-sae of this worm is large and reaches inwards
across the two longitudinal tubes of the water-vascular system.
It is perhaps half a millimetre in length. As already mentioned,
the cirrus sacs alternate in position with absolute regularity in
successive proglottids. Each cirrus-sac opens on to the exterior
in mature segments rather behind the middle of the proglottid,
and its position is oblique—the posterior end being directed
anteriorly; the external aperture is thus directed vather back-
wards. I observed no individuals im copula, and it is dificult to
understand how this takes place. In the case of other Acoleide,
which are all, like the present genus, without female orifices in
the mature seoments, it has been suggested that the spiny cirri
perforate the body. Of any such spines there seems to be no
trace in the present species. But frequently the cirrus within
the sae was dilated with masses of sperm, conspicuous on account
of its deeply staining with hematoxylin. The cirrus-sac projects
into a fairly deep genital cloaca which is, in the most anterior
segments where the cirrus-sac is immature, quite as deep as the
cirrus-sac is long. The distinctiveness of the genital cloaca is
rather lost in the mature proglottids. The cirrus-sac itself is
oval in form, with a gradually decreasing anterior region which
projects into the genital cloaca. It is, in fact, pear- shaped with
rather a long stalk.
When the cirrus-sac is quite fully formed, it is seen to le in
a rather specialised region of the body parenchyma. This region
is formed of very lax tissue, which might thus be supposed to
allow of greater freedom on the part of the cirrus-sac. More-
over, isolated but numerous muscular fibres run inside and outside
of this lax area which are attached to the cirrus-sac and probably
serve as retractors. This lax area is not limited to the region
occupied by the cirrus-sac alone; it runs back and accompanies
the first part of the sperm-duct. In a series of sections the
cirrus-sac and the first part of the sperm-duct are seen to he
loosely in the lax tissue which extends beyond it in every
direction. The cirrus-sac is extremely muscular, the walls being
unusually thick. The muscle-layers are two, the cutside being
of fibres having a longitudinal direction, and within this is a
1046 - DR. F. E, BEDDARD ON
much thicker layer of circular fibres.. The inner layer is twice,
or in parts thrice, the thickness of the outer layer. The imner
circular layer ceases with the diminution in diameter of the
cirrus-sac on its way to the exterior; the inner oval region is
alone thus fortified. The outer muscular layer, however, con-
tinues to the distal extremity of the cirrus-sac and its narrower
projection. Outside of these muscle-layers the cirrus-sac is
enclosed within a single layer of rather large vesicular cells with
a prominent nucleus. It appeared to me that this layer was
continuous with the epithelium of the ensuing vas deferens,
though special to the cirrus-sae in its peculiar structure. The
cirrus-sac, as usual among the Cestodes, contains delicate muscular
fibres and nuclei within it in addition to the cirrus. In the
younger stages in the development of the cirrus-sac the sac has
Text-figure 4.
Moneecocestus erethizontis.
A proglottid viewed in its entirety, to show the position and shape of the
ovary (ov.) and yitelline gland (v.g.).
ci. Cirrus-sac. w.v. Water-vascular tpbe.
more delicate walls in the rounded part of it, while the region
leading to the genital cloaca has thicker muscular walls; the
precise reverse occurs in the adult sac. The cirrus itself lies in
a perfectly straight line in the adult cirrus-sac, and is of even
calibre throughout when not distended with sperm; there is
nothing in the nature of a vesicula seminalis within the cirrus-
sac such as has been described in many forms. But when
charged with sperm, dilatations are formed locally which are
therefore to be regarded merely as local turgescences due to the
enclosed sperm, and not, as representing a definite dilatation
on the course of the duct, such as is the receptaculum upon
the course of the vagina in this and other tapeworms. In
A NEW MAMMALIAN CESTODE. 1047
the younger and more spherical cirrus-sacs of earlier segments
the cirrus is coiled. It perforates the muscle-layers of the
cirrus-sac to become continuous with the vas deferens without
any change of diameter.
The ovary of this worm is ventral in position, as in Shipleya
inermis. In horizontal sections it forms with the vitelline gland
almost a complete ring, since the ovary is semicircular and the
smaller vitelline gland serves to close the semicircle posteriorly.
The ovary is larger than the vitelline gland, and lies, of course,
more anteriorly in the segment. In transverse sections the ovary
is Seen to possess a longitudinal and tubular form, the greater
part of the cavity being empty. This is illustrated in text-fig. 2
(p. 1041). It is there seen to be pressed closely against the
transverse muscular layer bounding the medulla, and thus to lie
below the receptaculum seminis and also, to some extent, the
vitelline gland. The remarkable tubular form of the ovary in
young proglottids might lead to its being confused with the
commencing uterus, which, however, lies above it and on a level
with the receptaculum ovorum. The ovary has thus a flattened
form when viewed in its entirety. It is near to the middle of the
proglottid verging towards the pore side. The young cells, which
will become ova, are chiefly massed at the two ends right and left
of the tubular ovary, and this region, as shown in the figure
referred to, is somewhat dilated on both sides, forming an oval
sac. The course of the tube when viewed in transverse sections
is quite straight from side to side.
§ Vagina.
A careful inspection of the horizontal sections shows that,
although no vagina opens into the so far isolated receptaculum
seminis, the equivalent of a vagina would appear to be present.
I cannot otherwise interpret a narrow straight duct which opens
on to the exterior beside, and in front of, the cirrus-sac. This
duct passes towards the interior of the body, up to a point ona
level with the end of the outer half of the cirrus-sac; it is
therefore of very. limited extent. It ends at this spot in a
dilatation, an oval sac. I have seen this tube ending blindly in
a sac in four or five segments. There is thus no question of its
normal presence; but I have seen it in the more anterior seg-
ments only, but which are nevertheless well provided with gonads,
and a cirrus-sac as large as it is in the more posteriorly situated
segments. The slender character of the duct and the delicate
chamber into which it expands, remind me greatly of the conditions
obtaining in my genus Diplopylidium *. But in the latter worm
* P.Z.S. 1913, p. 562. The illustration depicting the vagina of Diplopylidium
(text-fig. 92, p. 563) may be compared with my description of the present species.
In the text of that paper I have remarked that only in Tetrabothrius is this reversed
position of pores. ‘This is an obvious lapsus calami for Tetradiscotyla.
1048 DR. F, E. BEDDARD ON
the small receptaculum passes posteriorly into a duct leading to
the internal portion of the female apparatus. It seems to me
to be impossible not to recognise in this tube a rudimentary
vagina; but the dilatation can hardly be compared to a recep-
taculum, since that exists elsewhere.
In the first few proglottids of the worm, the female efferent
apparatus is still more highly developed than in those proglottids
which have just been considered. I find, in fact, in the first six
or so of those proglottids in which the reproductive apparatus
generally is recognisable in its details, that the vagina (text-fig. 5)
Text-figure 5.,
Moncecocestus ercthizontis.
Part of a horizontal section through an anterior proglottid where the
vagina (v.) has not commenced to disappear.
g-c. Genital cloaca.
is a wide thick-walled tube which opens on to the exterior close
to, and in front of, the cirrus-sac on the cne hand, and communi-
cates with the receptaculum seminis on the other, thus forming a
continuous female efferent apparatus, like that of more normal
Cestodes. It is, perhaps, during this transient state that fertili-
zation is effected, also in the normal way vid this tube and not
through the interstices of the body parenchyma. I have noted
certain stages in the degeneration of the vagina; that part of it
which opens into the receptaculum seminis persists, after it has
A NEW MAMMALIAN CESTODE. 1049
ceased to be a pervious duct, as a narrow tract of tissue blocked
entirely by medullary parenchyma, but still retaining a definite
wall (text-fig. 6) separating it from the surrounding parenchyma.
As already mentioned, the external part of the duct persists longer
as a pervious tube ending blindly and in a dilatation. This worm,
therefore, is interesting as showing a transition between such a
family as the Anoplocephalidz and the Acoleide in point of its
female efferent apparatus.
Text-figure 6.
a
iatiean
nee oo
:
Ny a ae
Pee
“>
ca
Monecocestus erethizontis.
Part of a horizontal section through a proglottid a little further back than that
referred to in text-fig. 5, to show degeneration of vagina (v.).
r.s. Receptaculum seminis. ¢. Testes.
The uterus of this worm is reticulate, except in very old
proglottids in which a more complete fusion of the network tends
to produce a saccular uterus, There are, however, even here,
traces of the reticular condition. These may ultimately perhaps
disappear. The uterus appears early in the body, and I have
depicted it in young proglottids in text-figs. 7 & 8, which are re-
spectively horizontal and transverse sections through young
segments. A corresponding section (text-fig. 9) represents the
mature uterus in subsequent segments. ‘This organ is seen to
lie quite medianly in the proglottid, that is to say, it is well
above the ovary and in the same straight line with the receptac-
ulum seminis. The ovary has disappeared in the more mature
proglottids. The uterus at first forms a network of rather
delicate strands which are abundantly nucleated but contain no
lumen, or only a very narrow one. In transverse sections the
lumen is quite visible in young proglottids, which are, however,
1050 . DR. F..E, BEDDARD ON.
rather older than that depicted in text-fig. 8. The cavities appear
in such a section to be detached, as the network is not so close as
it becomes later. The cavities are lined with an epithelium which
is at least less obvious in older uteri.
Text-figure 7.
Moncecocestus erethizontis.
Horizontal section illustrating the first appearance of the uterus (wt.).
7.s. Receptaculum seminis. w.v. Water-vascular tube.
In all the sections referred to it may be seen that the uterus
does not extend laterally beyond the trunks of the water-vascular
system. The uterus is quite persistent, and is not replaced by
anything of the nature of paruterine organs. The ripe eggs
have a delicate shell, as appears to be the case in the Acoleide
generally. L ee at
Naturally I have endeavoured to ascertain how far my species
A NEW MAMMALIAN CESTODE, 1051
resembles Bertia (now written Bertiella) americana, a species de-
scribed by Stiles * from the same genus Hrethizon, and previously
assigned by the American helminthologist to the genus Andrya.
This species, which is possibly, according to Stiles, identical
with Tenia laticephala of Leidy*t, is regarded by v. Janickit
to be identical with his Schizotenia.
Text-figure 8.
Monceecocestus erethizontis.
Transverse section showing the retiform uterus (wt.).
ci. Cirrus-sac. w.v. Water-vascular tube.
At first sight of the figures given by Stiles in explanation of
this species, I was disposed to think that it might be identical
with that which I describe in the present paper. For example,
the male pores alternate with absolute regularity from right to
left in segment to segment, and the very early development of
* Proc. U.S. Nat. Mus. xix. 1897, p. 165.
+ Proc. Ac. Sci. Philad. 1855, p. 443.
t Zool. Anz. xxvii. 1904, p. 782.
1052 _ DR. F. E, BEDDARD ON
the sexual organs is as in my species. The drawing * showing this
alternation does not clearly represent the female ducts; but they
are shown in a figure of a transverse section T. | infer them to
be present in “ Bertia americana,” though no particular and de-
tailed mention of them is made in the text. The shape of the
uterus, however, and the extreme narrowness of the segments,
are sufficient to prevent any confusion between the two species.
Tenia pectinata, described by Cobbold = from the same animal, is
Text-figure 9.
Monecocestus erethizontis.
Horizontal section showing the retiform uterus (wt.).
r.s. Receptaculum seminis. w.v. Water-vascular vessel.
too imperfectly known to permit of a comparison ; moreover, it
has unilateral genital pores, which are fatal to such a comparison.
The question may arise—are not my specimens perhaps
abnormal ?@ Von Janicki§$ has lately directed attention to two
species of Hymenolepis in which the genital pores are absent ;
and at the same time Wolffhiigel || has made similar observations
upon a species of Bertiella. These facts are applied by v. Janicki
* Stiles, loc. cit. pl. x. fig. 7. + Stiles, Joc. cit. pl. x. fig. 9.
- t Can. Nat. & Geol. vii. 1862, p. 394. ; hey
§ CB. Bakt. u. Paras. xxxvi. 1904, p. 222.
|| Berliner tierairztl. Wochenschr. 1904 (quoted by v. Janicki and Fuhrmann).
A NEW MAMMALIAN CESTODE. 1053
to Fuhrmann’s genus Aporina, among the characters of which
genus is the lack of these pores. Fuhrmann, however, has
pointed out * that even if in the future examples of Aporina alba
be met with in which the sexual ducts reach the exterior, the genus
Aporina will not be invalidated thereby. I may use the same
argument with regard to the species of Cestode which I describe
in the present paper. But if I were compelled to ignore the
rudimentary vagina, the genus would have to be placed in the
subtamily ‘Anoplocephaling of the family <Anoplocephalide,
though it would remain distinct.
Comparison with Shipleya inermis Fuhrmann.
Although at first sight, and indeed after some study, I was
disposed to place the present species in the same genus as
Shipleya, and even to consider the possibility of the iwe forms
being identical, Iam no longer able, after a more profound study of
this species, to identify even generically the two forms referred to.
It is remarkable, however, that in pursuing my supposed new
species through the dichotomous table of Mr. Ransom I arrived
at the genus Shipleya, the only difference apparently being the
Jack of spines upon the cirrus in my species. A reference to
Fuhrmann’s original description of Shipleya inermis shows, bow-
ever, that there are other differences, and of importance. Shipleya,
like other Acoleide, has a body-wall which is traversed by a
complicated series of muscles. Outside of the inner transverse
muscles is a double row of large bundles; outside of this again
are four separate and very thin layers of transverse fibres,
between each of which is a layer of longitudinal fibres. Such a
section is figured by Fuhrmann. In my species, on the other
hand, there is but a single and the usual layer of transverse
fibres, outside of which is a layer of singly disposed longitudinal
fibres, an arrangement in marked contrast to that of Shiples ya.
In the second place, although my species from Hrethizon has
a large cirrus-sac, this organ is not so large as in Shipleya,
where it measures no less than one millimetre in length and
is, furthermore, covered by diagonal fibres. In my species
the fibres have the arrangement described above on p. 1045.
There is, however, in both species the accurate alternation in
the position of the genital apertures. But we shall see directly
that another tapeworm, more probably allied to my species,
shows the same regular alternation of these pores from right to
left side of the body. I do not think it likely that I have failed
to see the spines upon the cirrus if they were really present and
like those of Shipleya; for Fuhrmann described these spines as
arranged in three rows and very like the thorny spines of the
Acanthocephala. I have seen nothing of the kind, though the
cuticle covering the organ is composed of almost separate rod-
like spinules. These, however, are closely set and in contact,
* Zool. Jahrb., Suppl.-Bd. x. 1908, p. 39.
1054 DR. F. E. BEDDARD ON
and cannot be compared with spines such as I infer to exist in
Shipleya. The last-mentioned genus does not appear to possess
testes in the same segments as those which contain the ripe
female sexual products; Fuhrmann did not see those gonads
but presumed that they were to be found in earlier segments.
Now in my species the testes are obvious and numerous in
very many segments. Prof. Fuhrmann, with his experience
of the structure of Cestodes, could not have missed them were
they so plentifully present in Shipleya mermis. This is a
most important difference between the two forms, and one
which marks out the genus Shipleya as having retained to
some extent the dicecious nature of its ally Diwococestus. There
is no trace of such a state of affairs in my species. It is doubt-
less of minor importance to point out that the receptaculum in
Shipleya has a crenate outline and isa small sac, while in my
species it is rather large and of circular contour. Also the
vitelline gland of my species is displayed in the same horizontal
section with the ovary, and therefore does not lie entirely dorsal
to it, as is stated to be the case in Shipleya. Finally, it is rather
remarkable that there should be so close a likeness between the
uterus in the two forms. It has in both a nearly annular shape,
being incomplete however on one side. The uterus persists in
both species, and is not replaced by anything in the nature of a
paruterine organ. This fact, coupled with the character of the
female generative system, leads me to place my species in the
neighbourhood of this genus Shipleya, but other details of
anatomy forbid their reference to the same genus.
The species described in this paper therefore differs from
Shipleya in the following assemblage of characters :—
(1) Lhe muscular layers of the body-wall are feeble.
(2) There are no papille on the scolex and no apical depression.
(3) The water-vascular tubes have no ramifications.
(4) The testes are nuinerous in all segments until those in which
the uterus is developed, and form rows right across the
proglottids.
(5) Although the vagina comes to be aborted it is fully developed
in the most anterior segments, and there are traces of the
terminal part for some way back in the shape of a sac
opening on to the exterior in front of the cirrus.
(6) The cirrus has no spines wpon it.
(7) The uterus forms a network.
(8) The vas deferens is dilated into a vesicula seminalis of
peculiar form.
As the definitions of genera among the Cestoidea go, these
characters are, as it appears to me, quite sufficient to allow of
generic separation. They are also accompanied by a few minor
differences, such as the form of the receptaculum seminis, and
also by some minor points of likeness, such as size and absence of
neck and regular alternation in genital pores.
A NEW MAMMALIAN CESTODE. 1055
The following characters will define the new genus, and the
name will suggest its affinity with the Acoleide :—
MONGCOCESTUS, gen. nov.
Scolex unarmed; proglottids not longer than broad. Genital
pores regularly alternating. Water-vascular tubes two pairs lying
side by side, connected by transverse vessel from inner tubes in each
proglottid; no network. Longitudinal muscles feebly developed,
without bundles. Generative organs visible in first or second pro-
glottid ; first genital pore in segment 6. Testes numerous im
transverse rows posteriorly, within area bownded by water-vessels ;
sperm-duct at first very wide and covered by glandular cells, after
this short and narrow, without coil or vesicula seminalis; cirrus-
sac large and very muscular, cirrus unarmed. Ovary curved in
front of smaller vitelline gland. Vagina present in a few early
segments, later aborted, with exception of spherical receptaculum
seminis. Uterus retiform, meshwork later tending to confluence.
Eggs with delicate shell.
For the present the above definition will have also to serve
for the new species, which I call MWonecocestus erethizontis, sp. n.
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THE SECRETARY ON ADDITIONS TO THE MENAGERIE, 1057
EXHIBITIONS AND NOTICES.
October 27, 1914.
Prof, EK. A, Mincuin, M.A., F.R.S., Vice-President,
in the Chair.
The Srcrerary read the following report on the Additions
made to the Society’s Menagerie during the months of June,
July, August, and September, 1914 :—
JUNE.
The registered additions to the Society’s Menagerie during the
month of June were 177 in number. Of these 84 were acquired
by presentation, 42 by purchase, 15 were received on deposit,
1 in exchange, and 35 were born in the Gardens.
The number of departures during the same period, by death
and removals, was 177.
Amongst the additions special attention may be directed to :-—
1 Anoa (Anoa depressicornis) 3, from Celebes, presented by
Willoughby Smith on June 4th.
1 Blesbok (Damaliscus albifrons), born in the Menagerie on
June 18th.
3 Gemsbok (Orywv gazella) § 3 2, from the Kalahari, new to the
Collection, purchased on June 2nd and 3rd.
1 Panolia Deer (Cervus eldi) 3, from Rangoon, presented by
H. L. Holman Hunt, F.Z.S., on June 13th.
1 Vicuha (Lama vicugna) 2, from Peru, presented by G. Noel
Clarke, F.Z.S., on June 12th.
1 Peruvian Tree-Poreupine (Coendw bicolor), from Peru, new
to the Collection, deposited on June 3rd.
2 Firecaps (Cephalopyrus flammiceps) and 1 Velvet-fronted
Nuthatch (Sitta frontalis), from India, both new to the Collection,
presented by Alfred Hzra, F.Z.S., on June 12th and 14th.
2 Maximilian’s Parrots (Pionus mawximiliana), from Brazil,
presented by the Marquess of Tavistock, F.Z.8., on June 18th.
2 Cotton Teal (Vettopus coromandelianus), from India, pre-
sented by Hubert D. Astley, F.Z.S., on June 29th.
1 Rufous-crowned Ground-Cuckoo (Cowa ruficeps), from®Mada-
gascar, new to the Collection, purchased on June 9th,
1 Arizona Snake (Coluber arizonce), from N. America, new to
the Collection, purchased on June 29th.
JULY,
The registered additions to the Society's Menagerie during the
month of July were 277 in number. Of these 93 were acquired
by presentation, 81 by purchase, 60 were received on deposit,
9 in exchange, and 34 were born in the Gardens.
Proc. Zoou. Soc.—1914, No. LX XI. fal
1058 THE SECRETARY ON ADDITIONS TO THE MENAGERIE.
‘The number of departures during the same period, by death
and removals, was 239.
Amongst the additions special attention may be directed to :—
1 Lesser Kudu (Strepsiceros imberbis) $, from Berbera,
presented by Arnold Hodson on July 28th.
1 Cape Ant-bear (Orycteropus capensis), from §$. Africa,
purchased on July Ist.
2 Canadian Beavers (Castor canadensis), born in the Menagerie
on July Ist.
1 Sanday Island Vole (Microtus sandayensis), from Sanday
Island, N. Orkneys, new to the Collection, presented by W. R.
Ogilvie-Grant, F.Z.8., on July 11th.
1 Black-necked Stork (Xenorhynchus asiaticus), from India,
purchased on July 28th.
1 Great Courlan (Aramus giganteus), from Cuba, new to the
Collection, purchased on July 4th.
1 Swainson’s Hawk (Gampsonyx swainsoni), from 8. America,
new to the Collection, purchased on July 27th.
1 Australian Ground-Thrush (Oreocinela lunulata). from New
South Wales, new to the Collection, purchased on July Ist.
1 Blue-headed Rock-Thrush (Petrophila cinclorhyncha), from
the Himalayas, new to the Collection, presented by Alfred Ezra,
F.Z.S., on July 27th.
1 Flame-breasted Robin (Petraca phanicea), from New
South Wales, new to the Collection, presented by Alfred Ezra,
F.Z.S., on July Ist.
1 White-fronted Chat (Zphthianura albifrons), from New
South Wales, new to the Collection, purchased on July Ist.
2 Scarlet-breasted Robins (Petraca leggi), from New South
Wales, new to the Collection, deposited on July 17th.
2 White-shouldered Caterpillar-eaters (Lalage tricolor), from
New South Wales, new to the Collection, presented by Alfred Ezra,
F.Z.8S., on July Ist.
5 Malachite Sun-birds, 2 g and 3 Q (Nectarinia famosa), from
South Africa, new to the Collection, deposited on July 8th.
4 Lesser Double-collared Sun-birds (Cinnyris chalybeus), 2 3,
2 9, from South Africa, new to the Collection, deposited on
July 8th.
_ 1 Friar-bird (Zropidorhynchus corniculatus), from New South
Wales, new to the Collection, purchased on July Ist.
2 Banana-Quits (Certhiola flaveola), fvom the West Indies, new
to the Collection, deposited on July 8th.
1 Collared Lizard (Crotaphytus collaris), from Western United
States, new to the Collection, purchased on July 4th.
1 Giant Monitor (Varanus giganteus), from Australia, new to
the Collection, deposited on July Ist.
2 Striolated Skinks (Lyernia striolata), from Australia, new to
the Collection, purchased on July 4th.
THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 1059
AUGUST.
The registered additions to the Society’s Menagerie during the
month of August were 200 in number. Of these 120 were
acquired by presentation, 32 were received on deposit, 17 in
exchange, and 31 were born in the Gardens.
The number of departures during the same period, by death
and removals, was 224.
Amongst the additions special attention may be directed to :—
1 Kordofan Giraffe (Giraffa camelopardalis) 3, born in the
Menagerie on August 12th.
4 New Zealand Geckos (Hoplodactylus pacificus), 3 Moco Skinks
(Lygosoma moco), from New Zealand, and 1 Slender Skink (Lygo-
soma tenwe), from Australia, all new to the Collection, deposited
on August 6th.
1 Smoky Snake (Tropidonotus fuliginoides) and 1 Burrowing
Viper (Atractaspis irregularis), from Sierra Leone, both new to
the Collection, presented by Guy Aylmer, F.Z.S., on August 6th.
SEPTEMBER. °
The registered additions to the Society’s Menagerie during the
month of September were 130 in number. Of these 56 were
acquired by presentation, 51 by purchase, 11 were received on
deposit, 6 in exchange, and 6 were born in the Gardens.
The number of departures during the same period, by death
and removals, was 161.
Amongst the additions special attention may be directed to :—
1 White-bearded Gnu (Connochetes albojubatus), born in the
Menagerie on September 28th.
2 Grey-tailed Fruit-Pigeons (Osmotreron griseicauda), from
Java, new to the Collection, presented by the Marquess of
Tavistock, F.Z.8., on September 18th.
A Collection of Birds from New Guinea, the Aru Islands, ete.,
acquired by purchase on September 11th, and containing Greater
Birds-of-Paradise (Paradisea apoda), a Lesser Bird-of-Paradise
(P. minor), King Birds-of-Paradise (Cicinnurus regius), and the
following species new to the Collection :—
1 Fairy Bluebird (/rena turcosa) from Java, 2 Blue-eyed
Ravens (Jacrocorax fuscicapillus), 1 New Guinea Pitta (Pitta
nove-guinee), and 3 Duperrey’s Megapodes (Meyapodius du-
perreyi) from the Aru Islands; and 2 Pied Kgrets (Wotophoy«
flavirostris) from South-west New Guinea.
Mr. R. H. Burne, M.A., V.P.Z.S., exhibited a number of
preparations showing some adaptations for the nourishment of
the embryos of Elasmobranchs.
71%
1060 THE SECRETARY ON ADDITIONS TO THE MENAGERIE.
Mr. R. E. Savage exhibited two abnormal Herrings (Clupea
harengus), taken by trawl in the North Sea. The first specimen
had neither pelvic fins nor girdle (pelvic bones). The usual
position of base of fins was indicated externally by the presence
of the characteristic elongated scales. The usual musculature
was present internally. The second specimen lacked the left
pelvic fin and pelvic bone; the musculature was complete.
Messrs. E. Heron-Auten, F.L.S., F.Z.S.,and Arraur Ear.ann,
F.R.M.S., read a paper on the Foraminifera of the Kerimba
Archipelago, obtained by Dr. J. J. Simpson in the years 1907-8.
This paper has been published in the ‘ Transactions.’
Mr. W. L. Distant communicated his Report on the Rhyn-
chota collected by the Wollaston Expedition im Dutch New
Guinea.
This paper has been published in the ‘ Transactions.’
November 10, 1914.
Prof. E. W. MacBrarpz, M.A., D.Sc., F.R.S., Vice-President,
in the Chair.
The Srcrerary read the following report on the additions to
the Society’s Menagerie during the month of October 1914 :—
The number of registered additions to the Society’s Menagerie
during the month of October was 290. Of these 268 were
acquired by presentation, 15 were received on deposit, 6 in
exchange, and 1 was born in the Gardens.
The number of departures during the same period, by death
and removals, was 207.
Amongst the additions special attention may be directed to :—
A Collection of Birds presented by Sir Walter Gilbey, Bart.,
F.Z.8., on October 12th, consisting of a Hunstein’s Bird-of-
Paradise (Diphyllodes hunsteini) trom South-east New Guinea,
a Hildebrandt’s Starling (Spreo hildebrandti) from Kast Africa
(new to the Collection), a Hunting Crow (Cissa chinensis) from
India, and a number of other rare birds.
A Collection of Birds from Chili, presented by Geo. H. F.
Duncan, F.Z.S., on October 22nd, containing 10 Cayenne Lap-
wings (Belonopterus cayennensis), 2 Chilian Sea-Hagles (Gera-
noaétus melanolewcus), 5 Gay’s Finches (Phrygilus gayi), and
others.
BIRTH OF A PORPOISE AT BRIGHTON AQUARIUM. 1061
2 Red-necked Phalaropes (Phalaropus hyperboreus), from
Iceland, new to the Collection, presented by W. H. St. Quintin,
F.Z.8., on October 8th.
The Secretary exhibited a photograh showing Oysters
growing upon mangroves at Lobito Bay, Portuguese West
Africa.
Mr. R. H. Burne, M.A., V.P.Z.S., exhibited some paraffin
Simulacra of Molluscan and other Shells made accidentally by
Dr. C. V. Ariens Kappers while embedding objects for the
microtome. During this process, paraffin in a molten state
inadvertently escaped from the mould and set in the shell-forms
shown, probably owing to distorted crystallization.
Mr. H. R. Hoce, M.A., F.Z.S., read his Report on the Spiders
collected by the British Ornithologists’ Union and Wollaston
Expeditions in Dutch New Guinea.
This paper will be published in the ‘ Transactions.’
Lirth of a Porpoise at the Brighton Aquarium.
Mr. Lewis H. James, B.A., F.Z.8., communicated the following
notes upon the birth of a porpoise at the Brighton Aquarium :—
On the 26th May, 1914, a female porpoise (Phocena communis)
was caught in a kettle-net at Dungeness. She was transferred
to a sluice immediately she was landed and left there for a short
time. She was then seen to be inanadvanced stage of pregnancy.
Later, she was transferred by road and rail to Brighton Aquarium,
where she was placed in a tank containing 110,000 gallons of
sea-water. During the journey the porpoise was wrapped in
wet blankets and the eyes and nares were frequently sponged
with sea-water. On being placed in the tank, she seemed in good
condition and swam well.
On the afternoon of Sunday, 31st May, the porpoise seemed to
be in great pain. She swam to and fro in the tank, resting at
intervals on the surface. This was about 3.15 p.m., and from
this time typical labour movements set in. While lying at the
surface of the tank she made straining movements by arching
her back, and in about half an hour the first signs of the offspring
appeared in the form of the tail. Labour was now very strained,
and the mother appeared in great pain, probably owing to the
fact of a breach presentation. About one-half of the young was
forced out by spasmodic contractions, while presentation was
completed by a final forced contraction. The whole process
lasted from 3.15 to 5.50 P.M.
1062 MR. R. I. POCOCK ON
The cord was still attached to the mother, but was soon brqken
‘after she had swum 20 yards or so.
After the birth of the young, which was a still-born male, ihe
mother was much easier, and before long was swimming about
normally. After about five hours, the first signs of the after-
birth appeared and was suecessfully passed 44 ‘hours later. As
described to me by the superintendent who witnessed it, the after-
birth was ‘a chamois leather-like paunch” and weighed 13 lbs,
Unfortunately it was thrown away. There were slight straining-
movements during the passing of the placenta, but very little
bleeding.
The young weighed 7 lbs. approximately and measured 2 feet
2 inches. It was perfectly formed, and its skin was soft,
resembling a kid glove.
The mother died on Wednesday, 10th June—probably, if she
had been allowed to remain in the sluice for a time, the fetus
would have turned successfully, and both mother and young
would have been saved. On the journey between Lewes and
Brighton the mother passed a certain amount of foaming blood
through her nostrils, but when placed in the tank seemed to
be perfectly fit.
The mother and young are at present at Brighton to be mounted
for the local Museum.
Pine and Beech Martens.
(Text-figures 1-4.)
Mr. R. I. Pocock, F.R.S8., F.L.S., F.Z.S., Curator of Mammals,
gave an exhibition, illustrated by lantern-slides, to show some
newly noted points of difference between the Pine Marten (J/artes
martes) and the Beech Marten (Jartes foina) and remarked :—
“The cranial and dental differences between these two species,
pointed out by Mr. E. R. Alston in 1879 (P. Z. 8. 1879, p. 468),
are well known, and have been recently restated by Miller (Cat.
Mamm. Western Europe, 1912). Alston scarcely dealt with
external characters, and Miller, when deseribing J/. foina, dis-
missed them as follows: ‘‘ external form as in J/. martes, but fur
of less fine quality ; colour usually more greyish or drab than
in IW. martes and seldom with the rich brown tints of [that]
species, the throat patch never strongly tinged with yellow.”
From this it might be inferred that the only external differences
between the two species are found in the texture and colour of
the fur. That the differences in the structure of the ears and
feet I am about to describe were overlooked by Miller, must
presumably be attributed to his working with dried skins and
not with specimens preserved in alcohol. The material upon
PINE AND BEECH MARTENS. 1063
which my conclusions are based are two examples of I. martes
and one of MJ. foina, sent from France to the Gardens for
Mr. Thompson Seton. They were not quite adult, but practically
of the same size. They died upon the same day in September,
after being kept under precisely similar conditions, and were
preserved in alcohol for the investigation of external characters.
Ears.—All the Pine and Beech Martens that have been ex-
hibited within my recollection in the Gardens could be distin-
guished at once, when seen side by side, by the difference in the
Text-figure 1.
Pine Marten (Martes martes).
size of the ears (text-figs. 1 & 2). In the Pine Marten these
organs are relatively longer and wider and appear in consequence
to be more pricked, so that the animal looks more alert than the
Beech Marten. Moreover, the narrower ears in the latter niake
the intervening space on the crown wider, and the whole head has
a broader look than in the Pine Marten. The actual differences
in dimensions in the ears are shown in the subjoined table of
measurements (p. 1068). In details of structure the ears are
much alike, as might perhaps be expected. In both species there
1064 MR. R. I. POCOCK ON
is a very distinct lobe upon the “antihelix”*; but whereas in
M. foina this lobe is thick and fleshy, in W/. martes it is com-
pressed and scale-like. Finally, the integumental flap at the back
of the ear, like the pinna of the ear itself, is longer in J/. martes
than in J, foina (text-fig. 3).
It is probable that the length and breadth of the ears will be
found to vary locally in both MW. martes and IM. foina; but many
more measurements than have as yet been taken must be
recorded before the average difference in size of these organs
between the two species can be established.
Text-figure 2.
Beech Marten (Martes foina),
Feet.—From the appearance of living animals I supposed at
one time that J/. martes was a longer legged form than J/. foina;
but measurements of specimens in the flesh show that this is
not so. The feet are apparently of practically the same shape
in the two. The four main digits are separated by subequal
intervals and webbed, as in the Canide, practically to the level
of the proximal margins of the digital pads. These digits, more-
over, are very nearly symmetrical in the sense that the second
and fifth and the third and fourth toes are respectively almost
* The plica principalis of Boas, ‘ Ohrknorpel und ausserer Ohr,’ 1912.
PINE AND BEECH MARTENS. 1065
Text-figure 3.
A. Head of M. martes, with ear fully expanded.
1. Lobe on antihelix.
B. The same of UW. foina.
A}. Side view of ear of M. martes, with the posterior half folded forwards
to show the flap (f°) on the back of it.
B}. The same of UW. foina.
1066 MR. R. I. POCOCK ON
ona level. The area of the sole between the plantar and digital
pads is thickly hairy except for narrow naked streaks, visible
when the hairs are parted, running forwards from the plantar
pad towards the digital pads. The area behind or above the
plantar pad is also covered with hair, except that in the fore foot
there are two carpal pads, a larger (hypothenar) on the ulnar
side of the middle line and a much smaller (thenar) on the radial
side. These two carpal pads are separated by a considerable
space from the plantar pad and by a very narrow hair-covered
space from one another. The main portion of the plantar pad
of both fore and hind feet is nearly bilaterally symmetrical,
consisting of the three normal lobes corresponding to the four
principal digits, but behind the inner lobe corresponding to the
second digit there is an accessory lobe or pad corresponding to
the first digit. The digital pads and the lobes of the plantar
pads are concentrically striate, and the summits of the lobes of
the plantar pad on the hind foot are crowned with a cluster of
small close-set papillee *.
In the particulars described above the feet of the two species
are alike; but so far as the pads are concerned considerable
differences may be noticed. The pads of JZ. foina may be de-
scribed as normal in size; the digitals are broadly elliptical and
a little longer than and nearly twice as wide as those of I. martes.
a plantar and carpal pads also are larger and encroached upon
by hair to a much lesser extent in J/, foina than in J/. martes.
In the hind foot of J/. martes indeed the plantar pad may be
said to be reduced to four striate tubercles joined by narrow
strips of hairless, soft, unstriated integument. These differences
are Shown in the figures and in the subjoined table of measure-
ments (p. 1068). The smaller size of the pads in JS, martes makes
the digits appear to be longer than in J. foina, but there does
not seem to be actually any difference in this respect between the
two species (text-fig. 4).
The hair on the metatarsal area is much thicker and woollier
in texture in J. martes than in M/. foina.
Ido not suppose that the distinguishing points in the pads
above pointed out will be found to be absolutely constant in all
specimens of these two Martens from different localities and at
all seasons of the year; but I strongly suspect that closely corre-
sponding differences will prove to obtain between specimens
collected in the same countries or latitudes at the same season of
the year. It is possible that in both species the pads are more
overgrown with hai in northern than in southern individuals,
and in winter than in summer, These points have, I believe,
yet to be worked out in detail. But it was that possibility I
had in mind when stating that the examples I examined, all
probably Central European, died upon the same day and had
* Martens are prodigious jumpers, and the papillz on the plantar pads probably
per iate the likelihood of slipping during their leaping pursuit of Squirrels through
the trees.
PINE AND BEECH MARTENS. 1067
been previously kept for several weeks under precisely similar
conditions *.
Text-figure 4.
Cee
NON Zenda
My it \ bs aN He 4
NIU AY
. ay A
(as)
A. Fore foot of MW. martes. | 3B. Fore foot of I. foina.
A!, Hind foot of the same. | B!. Hind toot of the same.
* Miller’s statement (op. cit. p. 368) that the feet of IZ. martes are densely furred
throughout in winter is certainly not always, if it is sometimes, true. It is con-
tradicted by the skins of two Cumberland specimens, killed on Dee. 5th, and pre-
served in the British Museum. The pads in these are quit> obviously bare and not
even concealed, much less overgrown, by the fur of the sole.
1068 ON PINE AND BEECH MARTENS.
Subjoined are the principal measurements of examples of the
Pine and Beech Martens, elucidating the points above described.
M.martes., M. foina. |
| |
Length of head and body .................0:2e0+.+-....], 380 mm.| 3880 mm. |
5 tar without tubs... sss. 0. sscseocenen che MEL Od oar Lis ee
is hind Tepstromphocks.,-saanee essen ee rere Sle SI;
iS front leestrom\elbowse.sc-se+- nc. ese se ses ces || ee OMe LOT 5.
| o Car ibromMunNOeCM eaeetesscs- tho ceereeee es aes aL 3 Bivane a
| Wadthvotearitullyispreadinccss.ue-e sede scene exe eeeeene Se on na aes tt
Length of median lobe of plantar pad of fore foot...
|: Wadthiofidithow::. :\ivees cnet acs sccss eee ere ae
Width of pad excl. pollical lobe .....................44- 1
| Genephiotend: diciiall pains scrpssvesceneseeeemaneeeneees
Width of ditto RGaeenicegie tals st toe eehid oe eee ae
engthrotilarcercarpalpadiss:..crsn.ssssnaseese eee
| Width of ditto eairewese
Orc WsTO1rm Or
The dimensions in millimetres of four specimens published by
Miller may be compared with those set forth above :—
}
M. martes. M. foina.
Como. | Minorea. Switzerland. ces,
| Length of head and body ...... 470 430 453 403
| Sue Stallion: Aspe eee well O35 230 260 | 255
Ba mbindtoober. secu loas6 87 85 eee
“p CA Ae rae CEE Ss 42 45 34 | 39
Making the necessary allowances for errors in small respects
and for variations due to the “personal equation” of the col-
lectors, this table bears out on the whole the contention that the
ears are longer in J/. martes than in I. foina. The Swiss speci-
men of the latter, an adult male from St. Gallen, has remarkably
short ears, whereas the Cretan example, regarded by Miller as
representing a distinct species (JZ. bunites), has these organs a
little longer than the specimen of J/. foina measured by myself,
although, judging by the length of the hind foot, it was a smaller
animal *, Miller described it as a “young adult male.” It
must be remembered in this connection that the ears in young
mammals are always relatively longer than in the adults and
probably do not increase appreciably in length after the subadult
stage is reached. Clearly, however, the average length of the ear
in the three examples of J/. foina is much less than the average
length of the ear in the three examples of MW. martes, the
dimensions being approximately 37 and 43 respectively.”
* The shortness of the tail and of the head and body in my examples is probably
attributable to the shrinkage of specimens preserved in alcohol, which could not be
properly straightened.
PURPOSE AND INTELLIGENCE IN THE FORAMINIFERA. 1069
November 24, 1914.
Prof. E. A. Mincurn, M.A., F.R.S., Vice-President,
in the Chair,
Mr. E. T. Newron, F.R.S., F.Z.S., exhibited a series of bones
of animals showing indications of natural repair, and a number
of teeth of a female Sperm-Whale (Physeter macrocephalus).
Purpose and Intelligence in the Foraminifera.
Messrs. KE. Herron-Auuen, F.L.S., F.Z.S., and Arraur
Bartanp, F.R.M.S., discussed the Phenomena of “ Purpose ”
and ‘‘ Intelligence ” exhibited by the Arenaceous Foraminifera
in the construction of their tests. They observed that W. B.
Carpenter had stated in 1885 that the highest development of
function and behaviour of protoplasm in the Protozoa was to be
found in the study of the Foraminifera, and they exhibited slides
of porcellanous and hyaline species, calling attention to the fact
that of two groups of the same Zoological Order, constructed of
the same simple protoplasmic element and living under similar
conditions, one group secretes calcium carbonate and the other
strontium sulphate from the same sample of sea-water, for the
construction of their shells.
The behaviour of many species of Arenaceous Foraminifera in
constructing their tests reveals a development of “ purpose”
amounting to what in the Metazoa would be termed “ intelli-
gence.” his display of intelligence and purpose takes two forms:
(i) The exclusive selection of certain materials out of a prepon-
derating mass of other and more readily utilizable materials; and
(ii) the manner in which those materials are used. This latter
is subdivided under two heads—(a) to meet the requirements of
the organism in adapting itself to environmental circumstances,
and (6) to protect itself against the attacks of parasitic worms
and other enemies.
Lantern-slides were shown, and the actual specimens exhibited
under microscopes upon the table, to illustrate the following
phenomena :—1, The selection of magnetite by Haplophragmium
agglutinans and of topaz and garnet by Vernewilina polystropha,
for the construction and decoration (?) of their tests. 2. The
utilization of foraminiferal shells by certain abyssal worms, and
the method employed by Pectinaria auricoma in “ laying” frag-
ments of sponge-spicules of even length in the construction of
its tube. 38. Psammosphera parva Flint, which builds a mono-
thalamous shell of sand-grains round a long sponge-spicule,
which is used as a “catamaran” spar to buoy it up upon the
bottom ooze. 4. Psammosphera bowmani WHeron-Allen &
Earland, which constructs its shell of flakes of mica cemented
LOTTO EGG OF THE NEW GUINEA RIFLE-BIRD.
together at their edges to form a transparent polyhedral test.
5. Psammosphera rustica H.-A. & E., which constructs a
polyhedral test between long ‘‘ catamaran” spicules, filling the
interspaces with fragments of spicules of graduated length and
occasionally using a triaxial spicule to fill an awkward space or
angle. 6. Vouria harrisii H.-A. & E., which supports itself,
aperture uppermost, upon the mud by means of spicules pro-
jecting at the aboral end of the test. 7. Haliphysema
tumanowiezii Bowerbank, which protects its aperture from
parasitic worms with an investment of projecting spicules.
8. Crithionina pisum Goés, var. hispida Brady, 9. Paulina
jetireysti Carpenter, and 10. Hyperammina ramosa Brady,
which protect themselves with a close investment of spicules
all over their shells. 11. JJarsipella cylindrica Brady, which
protects its aperture with a crown of loosely aggregated spicules.
12. Marsipella spiralis H.-A. & E., which arranges the spicules
of which it is constructed in a left-handed spiral to give tensional
strength. 13. Technitella legumen Norman, which constructs its
shell of fragments of spicules in two layers, the inner layer being
laid at right angles to the outer. 14. Techunitella thompsoni
H.-A. & E., which selects from the environmental material for
the construction of its test nothing but the perforated plates of
an ophiurid or echinoderm.
In the opinion of the authors, ‘ purpose” and “ intelligence ’
are revealed by these phenomena. If they depended on surface-
tension, all the individuals in a dredging would exhibit the same
phenomena, and the theory of natural selection resulting m the
survival of the fittest is met by the reply that these selective
and purposive individuals constitute a marked minority in the
dredgings in which they are found.
An interesting discussion followed, in which, amongst others,
Sir H. H. Howorth, F.R.S., Sir E. Ray Lankester, F.R.S., and
the Secretary took part.
?
Egg of the New GuineaRifle-bird.
My. D. Seru-Suirn, F.Z.8., Curator of Birds, exhibited an egg
of the New Guinea Rifle-bird (Ptilorhis intercedens) which had
been laid in the Society’s Gardens in July last. The bird that laid
it belonged to Mr. HE. J. Brook, F.Z.S., who had kindly lent
it in the hope that it might pair with a male belonging to
the Society. While in Mr. Brook’s aviaries in Scotland, this
bird paired with a male of its species, and in 1911 and 1912
constructed a nest and laid eggs, but in both cases they were
infertile.
The birds were placed last spring in the Summer Aviary, where
they appeared to do well, and in July the female built a nest,
composed of dead leaves and dry vrass, in a bush, about five feet
from the ground, and laid two eggs. She sat well and was not
DENTAL VARIATIONS IN MAMMALIAN SKULLS, 1071
disturbed in any way, but about three weeks after the eggs were
laid they were apparently thrown out of the nest by the “sitting
bird, for they were found broken upon the ground. Neither
appeared to have been fertilized. The one least damaged was
saved,
This is the first instance of any species of Paradise-bird laying
in these Gardens, and the eggs laid in Mr. Brook’s aviaries were
the first ever laid in captivity. The bird that laid these eggs
was captured on the nest in New Guinea by Mr. Goodfellow, and
the nest and two eggs upon which she was sitting are now in the
British Museum.
The egg is very similar to that of the Greater Bird of Paradise
(Paradisea apoda), and, so far as we know, characteristic of the
whole Broun The ground-colour is cream, bold streaks of
reddish brown radiating from the larger end and overlaying
fainter splashes of pale grey.
Dentat Variations i im M ammalian Skulls.
Dr. Roperr Roan, G. M. Z. S., exhibited :—
(1) A number of skulls of Zrichosurus vulpecula illustrating
dental variations. Normally upper p’ is situated 2°5 mm. behind
the canine. Two skulls exhibited show it from *5 mm. to 1 mm.
behind the canine. One skull shows on the left side p' closely
pressed against the canine, and on the right side the two teeth
fused together. Another skull shows the p entirely absent.
In the lower jaw normally the small tooth behind the large
incisor is close to it. One jaw shown has the second tooth
2 mm. behind the large incisor. A second jaw has 2 mm.
behind the normally placed second tooth a third small tooth,
which may be either a third incisor, a canine, or a first
premolar.
(2) A skull of Phascolarctus cinereus showing in the right
lower jaw a small tooth behind the large incisor, resembling
in position the second tooth in the medinle of Tr falvecninne
vulpecula.
(3) A series of skulls of Chrysochloris hottentota and C. asiatica
illustrating the peculiar loss of teeth found in nearly all sexually
mature Moles in the Stellenbosch district,S. Africa. As Dr. Broom
pointed out in 1907, C. hottentota, when probably. two years old
and when in full sexual activity, suffers from a disease of the
gums, probabiy pyorrheea, and the teeth become loose and fall
out. The disease appears also to damage the developing second
set, which never fully replaces the lost milk set, and hence
specimens are found where the disease has been arrested in a
practically toothless condition.
It is very interesting to note that these toothless specimens
1072 DR. R. BROOM ON
appear to be in fairly good health. In their stomachs are the
remains of worms broken into segments, as in the fully toothed
individuals. °
Though specimens in a similar condition have been obtained
from other parts of S. Africa, they are rare.
Chrysochloris asiatica varely suffers with its jaws, but a speci-
men was exhibited in which the disease had led to extensive
injury to the bone.
(4) A skull of Chrysochloris hottentota with only eight teeth in
each upper jaw, though there are the normal nine on each side
of the lower.
A new Thecodont Reptile.
(Text-figures 1, 2.)
INDEX. Page
Eosucura, new suborder of THECODONTIA ......... 1077
SVOUNP INI ey LAIN. MOMs ay seeneneees<ceancsiedeese-enee rene TA
Youngina capensis, gen. et 8p. Ne .......seceeeescee ees 1075
Dr. Broom also exhibited the skull of a new type of Thecodont
reptile (Youngina capensis) from the Upper Permian beds of
South Africa, and remarked :—
“ Hitherto, though there has been some evidence of the occur-
rence of primitive Thecodonts in the Karroo beds of Upper
Permian age, no satisfactory skull has been known. Some years
ago I described the two types, Heleosaurus scholtzi and Heleo-
philus acutus, both of Middle Permian age, and Watson is
describing another form of the same age which may be allied.
Unfortunately, little is known of the skull in any of these types.
The new form which I ain exhibiting was found by me at New
Bethesda, Cape Colony, in beds which are in the Cistecephalus
zone, at least 600-1000 ft. below the Lystrosawrus zone, and most
probably belong to the Upper Permian age.
The specimen consists of the almost perfect skull of a small
crocodile-like reptile, with a considerable series of vertebra, but
unfortunately with no satisfactory remains of limbs or girdles,
The skull measures about 60 mm. in greatest length, and is -
42 mm. in width. The tip of the snout is missing, but as the
anterior ends of the lower jaws are preserved, very little can
have been lost.
The orbits, which are situated near the middle of the skull,
are large and are directed upwards and outwards. The antero-
posterior diameter of the orbitis 17mm. The interorbital width
is about 8 mm,
Behind the orbit is an infratemporal fossa slightly smaller
than the orbit, and a supratemporal fossa about a third of the
size of the infratemporal.
A NEW THECODONT REPTILE, 1073
The premaxillaries are lost but must have been small.
The nasals are imperfect and crushed; they are manifestly
fairly large. The nostrils must be situated well forward.
The maxilla is long and narrow, and on the right side shows
16 rounded pointed teeth. The complete series is probably about
21, a number evidently having been shed and were being replaced.
The posterior end of the maxilla passes under the orbit, and has
a long articulation with the jugal.
Text-figure 1.
Upper side of skull of Foungina capensis. About 13 times nat. size.
Fr., frontal; IP., interparietal; Ju., jugal; Na., nasal; Pa, parietal; Po., Par-
occipital: Po.F., posttrontal; Po.O., postorbital; Pr.., prefrontal; Q.J.,
quadratojugal ; So., supraoccipital ; Sq., squamosal ; Tb., tabular.
The lacrymal is not well preserved on either side, but is
manifestly small.
The prefrontal is a fair sized element. It forms the upper
and anterior border of the orbit, and, with the postfrontal,
nearly shuts out the frontal from the orbit.
The frontal is moderately large. It extends in front a little
beyond the plane of the front of the orbit, and behind as far as
the plane passing through the front of the pineal foramen.
Proc. Zoou, Soc.—1914, No. LX XII, 12
1074 DR. R. BROOM ON
The jugal is a long triradiating bone. Its anterior process ~
forms the lower border of the orbit, and passes forward between
the lacrymal and the maxilla. The ascending process passes up
behind the orbit to articulate with the postorbital ; while the
posterior process forms the lower border of the infratemporal
_ fossa, and lies above and internal to the quadratojugal,
The postfrontal is a small triangular bone which forms the
upper and anterior border of the orbit, and the anterior border
of the supratemporal fossa. It articulates with the frontal and
parietal internally, and the postorbital externally.
Text-figure 2.
————__—
eas So
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. ”
— B
A. Side view of skull of Youngina capensis. About 1} nat. size.
B. Occiput of Youngina capensis. About 1} nat. size.
IP., interparietal; Ju., jugal; La., lacrymal; Mx., maxilla: Pa., parietal;
Pa.O., paroccipital; Po.F., postfrontal: Po.O., postorbital; Pr.F., prefrontal ;
ne euereiee Q.J., quadratojugal; So., supraoccipital; Sq., squamosal ;
The postorbital is moderately large. It forms part of the
postorbital margin and most of the bar between the two
temporal fosse. The posterior process is well developed and
passes back between the squamosal and the tabular.
The parietals are smaller than the frontals. Between them is
a fairly large pineal foramen. The posterior process of the
A NEW THECODONT REPTILE. 1075
parietal is directed outwards and backwards, as in lizards and
Sphenodon, and rests on the bone which I believe to be the
tabular.
The structure of the post-temporal region is a little difficult to
make out, but as it is well preserved on ‘both sides the difficulty
consists largely in distinguishing cracks from sutures, and I
think there is practically no doubt but that the elements are
as I have figured them. Whether my interpretation of them
is correct is, of course, another matter.
Forming the posterior border of the infratemporal fossa is a
fair sized triangular bone, which [ believe to be the squamosal.
It articulates above with the postorbital, and below it probably
articulates with the quadratojugal. The quadrate is very largely
hidden by it.
With the posterior end of the jugal articulates an element
which I believe to be the quadratojugal. Unfortunately its -
posterior portion is not seen on either side. It cannot be a part
of the element which I identify as squamosal. The relations of
the quadratojugal, squamosal, and jugal are somewhat similar
to those in Sphenodon.
Along the upper and posterior side of the squamosal lies
another element of considerable size, which I believe to be the
tabular. In front it meets the postorbital, and its inner side is
partly covered by the posterior process of the parietal. Posteriorly
it articulates with the paroccipital (opisthotic).
Behind the parietals and above the supraoccipital is a fair sized
and certainly distinct interparietal. Possibly it is paired.
The occiput, though fairly well preserved, does not show the
limits of the various elements. The supraoccipital is low and
broad ; the exoccipitals are probably small, and there are short
paroccipitals directed backwards and outwards. Between the
sides of the supraoccipital and the tabulars are a pair of narrow
fosse. On the whole the resemblance of the occiput to that of
the lizard is considerable, the presence of a distinct interparietal
being the only important difference.
The mandibles, though present and complete, have not been
cleared of matrix, as it would be difticult to do so without
injuring the teeth, and the presence of the mandibles renders
it impossible to display the palate.
For this new reptilian type I propose the name Youngina
capensis, in honour of the late Mr. John Young, LL.D., F.G.S.,
Under-Curator of the Hunterian Museum, Glasgow University,
to whose early assistance and kindly encouragement I am mainly
indebted for my interest in paleontology.
Though Youngina cannot be placed close to any previously
well-known type, we can nevertheless give it a fairly definite
place in the reptilian series.
If we exclude the Cotylosauria, the Therapsida, the. Chelonia,
the Ichthyosauria, and the Plesiosauria, all the remaining orders
i
1076 DR. R. BROOM ON
. may be grouped together as reptiles with two temporal arches or
modifications of the same type. In Sphenodon we have a simple
modification of the type; in the Crocodile another, and in the
Lizards a third where the lower arch is lost. Among extinct
forms we have many other modifications. The Phytosaurs and
Pseudosuchians afford the best known Triassic types, and other
varieties of the type are seen in the Gnathodonts and the Dino-
saurs. Youngina represents a type more primitive than any
previously known, and one which is especially important in that
it is very near to the ancestral form.
The Pseudosuchian, Huparkeria, which I recently described
from the South African Upper Triassic beds, bears considerable
resemblance to Youngina but is very manifestly a much later
type. It has lost the large pineal foramen seen in Youngina,
and the post-temporal region differs considerably through the
loss of the tabular, the reduction in size of the squamosal, and
the increased development of the quadratojugal and quadrate.
Yet such a type as Yuparkeria might readily be descended from
a form like Youngina.
Euparkeria further represents a type from which the Thero-
podous Dinosaurs might be derived, but is too far advanced to
have been ancestral either to the Sauropoda or Predentata.
Youngina, on the other hand, retains the characters that we
require in the ancestor at least of the Sauropoda, and possibly .
also of the Predentata.
But the most interesting point in the structure of Youngina is
the light it throws on the origin of lizards.
No point in reptilian structure has given rise to so many
different opinions as the nature of the post-temporal region’ in
lizards. In Sphenodon there is no difficulty. There is low down
an undoubted quadratojugal, and between this and the parietal
a single large bone which is unquestionably the squamosal. In
the typical lizards, on the other hand, between the top of the
quadrate and the parietal are two small bones, and the difficulty
is to determine which is the squamosal. The upper and inner
bone has been regarded as the squamosal by Gegenbaur, Baur,
Gaupp, Case, and Watson; the lower and outer by Parker,
Huxley, Cope, Boulenger, and Williston. Until a year ago I
favoured the view of Baur, but the study of the Mosasaur skulls
in the American Museum led me to adopt the view of Williston,
that the outer bone is the squamosal and the inner the tabular.
About a dozen years ago I made a study of the development
of the pterygo-quadrate bar in a number of lizard types, and
found that the lower end of the quadrate is fixed to the lower
end of the epipterygoid by a small bar of cartilage almost exactly
as in Sphenodon. So strikingly similar is the condition that it
seems extremely probable that there was in the ancestral lizard
a lower bar as in Sphenodon.
While Youngina is certainly not a lizard, it throws very definite
A NEW THECODONT REPTILE: 1077
light on the nature of the lacertilian post-temporal bones. The
upper arch is formed mainly by the postorbital but partiy by a
lower bone which I regard as squamosal. Whatever this lower
bone is, it is quite certainly the homologue of the outer post-
temporal bone of the lizard, and as there is a quadratojugal still
lower down, it cannot be the quadratojugal. From its relations
to the quadrate, there seems little doubt but that it is correctly
identified as the squamosal.
The bone internal to it may be the so-called ‘ supra-temporal,”
but from its articulating with the paroccipital it seems much
more likely to be the tabular.
Youngina may be regarded as the type genus of a new family,
which may be called Younginide, and it must also be placed in a
new suborder of the Thecodontia, which may be called the
KosucHIA.
The Eosuchia may be defined as Thecodont reptiles which have
no antorbital vacuity, and retain the interparietal and tabular
bones, and have a large pineal foramen.”
2
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Exhibitions and Notices (continued).
Mr. D. Serx-Suitn, F.Z.S., Curator of Birds, Exhibition of an Egg of the New Guinea
Dr.
Dr.
ee
Rifle-bird (Ptilorhis intercedens) laid in the Society’s Gardens
Rozert Broou, C.M.Z.S. Exhibition of Mammalian Skulls showing Dental Variations.
Rosert Broom, C.M.Z,S. Exhibition of the Skull of a new Thecodont Reptile from
Dene AMIGA, wr ( hext-imuses! Lf Oi as)" e cae os ie wie « ss tine cee tls a stat oe = mistelon aie. eromie ole
PAPERS.
. A remarkable new Cirripede from the Chalk of Surrey and Hertfordshire. By ‘noms
Be warns, GS... (Plate I, and Dext-fcure 1.) «1.2. ees scseur wee coceseecuees
. Polycheta from the N.E. Pacific: The Chetopteride. With an Account of the
Phenomenon of Asexual Reproduction in Phyllochetopterus and the Description of
Two new Species of Chatopteride from the Atlantic. By F. A. Porrs, M.A., Fellow
of Trinity Hall, Cambridge, and Balfour Student of the University. (Plates I—-VL.,
sunaneret free UT ec Lehr)! 231 s)aeineiwia Sita -)afriveln 6 cstienje aS ¢ aie «ic oe eines weap amie Bele ne
, Broomia perplexa, gen. et sp. u., a Fossil Reptile from South Africa, By D. M. 8.
Warson, M.S8ce., F.Z.8., Lecturer on Vertebrate Paleontology in University College,
uname atate VL. sands Vext-truKesl—o.). © ja )sis)< chaielo cine) /cie:s)e ons sie oles aieieirs was
. Hunotosaurus africanus Seeley, and the Ancestry of the Chelonia. By D. M. S.
Watson, M.S8c., F.Z.S., Lecturer on Vertebrate Paleontology in University College,
uncon @alate. Vly ands Dextafroure Ws) oc.cisiicinlers a lels aie) eters cielo sieht s cnimisi sie cores «
. Notes on some Carnivorous Therapsids. By D. M. 8. Watson, M.Se., F.Z 8., Lecturer
on Vertebrate Palzeontology in University College, London. (Text-figures 1-7.)....
Contributions to the Anatomy and Systematic Arrangement of the Cestoidea.—
XV. On a new Genus and Species of the Family Acoleide. By Frank E.
Bepparp, M.A., D.Sc., F.R.S., F.Z.S., Prosector to the Society. (Text-figures 1-9.)..
EDEE Ves ee Ometateyeretteepeeetor she eich tatesreh cicte ok teaalSitiiny 0a is sae) sich qisyelay Sioasavcsnc aye Wwiallo) Saotetayous era MaE ates
Listiot. Councileand: Omicersss se oo eis sek es eee ee Heche tt grace as ey a encna, A, ar acai e d
Hist of Contentsiy oases. hese eR et gi a Bs it ean LGM Sy xt i ih a oan cpt Rein eevee ie
Alphabetical List of Contributors
2 G16 (bea, Me mrnars aia PU nay eg ese ROM cea a rs Set stl eM aap Pte female a. | BNnee LST Spe nemern sa Met day te tet
945
1011
1x
XV
LIST OF PLATES.
. 1914, Parr IV. (pp. 945-1077).
Wirners: Pl. I. 1-9. Proverruca vinculum, 10-12. Scalpellum
“ — edinineum wana wo Alls anaes ete aCe meee ;
Ports: Pl. I. Mesochetopterus taylori ....escecusvevcssees
II. Mesochetopterus minuta ......0ecseere sevoe
IIL, Mesochetopterus rh Satara lee een
es iy
V. \ Phyllochetopterus prolifica ...4..0e-ceeesseee
VI. Phyllochatopterus anglicd ssevresesesesenees
Warson: Pl. VI. Broomia perplexa .+...-sscecccesecceresceas
Warson: Pl. VII. Eunotosaurus africanus
NOTICE.
The ‘ Proceedings’ for the year are issued in four parts, paged ¢ co aseout iy
so that the complete reference is now P. Z. 8.1914, p. ..+ The Dice: D
is as follows :—
Part | I. issued in ‘March.
prea He ao a June,
pee (i a September,
pated th Aes ts December. ©
‘ Proceedings, 1914, Part ITI. (pp. 491-944), were published on n
September ath 1914.
The Abstracts of the ‘Proceedings,’ Nos.
: contained in this Part.
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