Cy PS ag EE,
f Me Z dl 4] ‘gt

}

Part 11. of Volume Il. will be published on
May 15, 1904.

The Fauna and Geography

of the
Maldive and Laccadive Archipelagoes

Being the Account of the Work carried on and
of the Collections made by an Expedition
during the years 1899 and 1900

Edited by

J. StanLey Garpiner, M.A.

Fellow of Gonville and Caius College and late Balfour Student
of the University of Cambridge.

VOLUME Il. PART Il.
With Plates XXXV—XLVIII and Text-Illustrations 120-126.

CAMBRIDGE }
at the University Press.

Lonpon: C, J. Cray anp Sons,
Cambridge University Press Warehouse,
Ave Maria Lane.

Price Fifteen Shillings net.

¢ ETLO900 TOEO g

UOMO A

1OHM/18IN

The Fauna and Geography
of the
Maldive and Laccadive Archipelagoes

VOLUME Il. PART Il.

Dondon: C. J. CLAY anp SONS,
CAMBRIDGE UNIVERSITY PRESS WAREHOUSE,
AVE MARIA LANE,

AND
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136, GOWER STREET, W.C.

Glasgow: 50, WELLINGTON STREET.
Leipsig: F. A. BROCKHAUS.
Pew Pork: THE MACMILLAN COMPANY.
Bombay and Calcutta: MACMILLAN AND CO., Lrp.

[All Rights reserved.]

The Fauna and Geography

of the
Maldive and Laccadive Archipelagoes

Being the Account of the Work carried on and
of the Collections made by an Expedition
during the years 1899 and 1900

Edited by

J. STANLEY GARDINER, M.A.

Fellow of Gonville and Caius College and late Balfour Student
of the University of Cambridge.

VOLUME Il. PART Il.

With Plates XXXV—XLVIII and Text-Illustrations 120—126.
+

CAMBRIDGE !
at the University Press.
1903

CAMBRIDGE:
PRINTED BY J. AND C. F. CLAY,

AT THE UNIVERSITY PRESS.

CONTHINGES, OF > VO die seAssr sels

Reports.

1. Marine Mollusca. With Plates XXXV and XXXVI
By Epear A. Smita, 1.8.0.

2. The Enteropneusta. With Plates XXXVII—XLVI and Text-
Figs. 120 and 121 , : d f : ; 7
By R. C. Punnert, M.A.

3. Marine Crustaceans. X. The Spider-Crabs (Oxyrhyncha). XI. On
the Classification and Genealogy of the Reptant Decapods.
With Plates XLVII and XLVIII and Text-Figs. 122—126

By L. A. Borrapatue, M.A.

PAGE

589

631

681

MARINE MOLLUSCA.
By Epear A. Smiry, I[.8.0.

(With Plates XXXV. and XXXVI.)

As the first collection of Marine Mollusca of any importance that has hitherto been
obtained at the Maldive Islands, that about to be described is of considerable interest. It
consists of about 380 different species and probably comprises a large proportion of the
forms which occur in these islands. That many have escaped notice and will eventually be
collected by others is quite certain, but it seems probable that most of the larger species
are represented in this collection and that it is only among the smaller and obscurer forms
that many additional species will hereafter be discovered.

\ There are, however, certain common and widely distributed species which one would
have expected to have been found, notably belonging to the genera Conus, Terebra, Sistrum,
Purpura, Nassa, Oliva, Solarium, Trochus and Circe. It is also worthy of remark that
certain genera which have representatives in the surrounding seas do not appear to have
been met with. Of these I may mention Fasciolaria, Rissoa, Patella, Eulima, Hydatina,
Aplysia, Umbraculum and Mytilus.

As might be expected the fauna is similar to that of other islands in the Indian
Ocean, many of the species occurring in Ceylon, Mauritius, Madagascar, the Amirantes,
Seychelles, and a great many ranging as far north as the Red Sea and Persian Gulf.
Rather more than a seventh of the forms have been recorded from the Andamans, and
probably many others occur at these and the neighbouring Nicobar group. It is well known
that great numbers of marine molluscs have a very wide range, and therefore it is not
surprising that many, in fact about three-fourths, of the Maldive shells are known to occur
in the seas surrounding the Philippine Islands and the Malay Archipelago, and still further
north about one-fourth of the species have been recorded from the islands of Japan. In
the Pacific, from the East coast of Australia to the Loyalty Islands and Polynesia, rather
over three-sevenths of the Maldive species have been recorded, many of them ranging all
over both the Indian Ocean and the Malay region also. About one-third of the species
are known from Ceylon, but doubtless many more occur there, although they may not have
been recorded. About 177 species have been found at the Mauritius, the fauna of which
has been more extensively worked than that of the Seychelles. With regard to the general
distribution of the species about to be recorded, it is curious to observe that a larger
proportion of them have previously been noted from the China Sea eastward and in the
Pacific, than in the Indian Ocean, including the Red Sea and Persian Gulf.

The following table shows at a glance the distribution of the species as far as the
writer has been able to ascertain without a very exhaustive search. The column under
Malaysia includes species which occur in the China Sea, the Philippine Islands, Malay
Archipelago, N. Australia, New Guinea, and as far east as the Solomon Islands. Under
Polynesia are indicated the species which are met with either in E. Australia or at the
Loyalty Islands, or at the various groups of islands of Polynesia.

G1. 76

EDGAR A. SMITH.

x sa BI 5 3
BP 2 1s \es) 8 | Sees
CRC aienieeme |e 0) ae
a ae | a 4 calc}
Solidula glabra....... ae
I nitidula .. + ais a ae
5 solidula . 3 Sl + + 4 +
7 LERSE LICL! © tcc eee tesa sree de
Bulla, GMpula wi ic...00.00ceeeaicnes «ose + + + ak st + +e ot
AlYJS CYUMATICG.. .c0.cc5c--n+sns0nn soon + aa + + a ds
of) EOERRGTIL one cooacooncooaapessonsesase: + Bs + uy ae
RR PSULC CUS Chucncins tied cci dono sis oitekriiran +
SWoluatellaecincta nn .ccer-eseer saree +
Oaynoe delicatula,.............000006 + + +
Cryptophthalmus minikoiensis, n. sp.
CONUSLANENALUS net--caactene des acace sere + + + + ss + + +
sath PLOBEULUOUS HctratnsGaanc sn sucaeeraes ao SF + + a + + 2
Wy) RCAUUS ck hacecrechine saci ces deseciocs +P + + + + - +
ay. GERI DES sop oaceeocodonansedcoee: + + + ne + “+
“5 COMSOMS .ececccenrereccrccnecerees +P
oh) GEEHOIES Sarnstan or Ganpacriboreatees + =i3 +
BS CULUTILOUS oat ccrcivencteec emesis ae + + + + + +
Apu ENIUUNEUS pec aatscavvsestewctee tenes + +
ny» GLALETEIUS: acces obnenaceenedRenebdoce + + + + + + + +
fy GIGI esceseabonesosansqoadssc ae 3F 3h se + =F + a +
A) OYLIEF Gaoscacddneacacsotocentasanoae + + oe ae +
rp TOS AES pe cotoodiaemaccoecstaussonos ar ae +f + ae ae + + se
3 uwidus..... Rutten sc geowirene seine + 3 + ot + + + + +
Bp RHA sooccsanasadeogoooanansececac + + ae 45 + + + +
A GMMMETILUTULITUILG ote ies oe oF + + + a5 +
>, mussatella ... ar + 38 + =r + + Te
+ penmnaceus ... + ae ae or
yy Quercinus + = + + + +
Oy) LUCERO OS Ben och scoeasecneeonedonn + ae
TIM TC CSSELLELELS Mrendecerwesc see aactnaats + 35 + ae ae ae ae + +
” CCRT C ears esse Ssc60onoRdonehosond ae ar ata oF ot sr ar + ar ae
Ah UAT Oreosee aceenchtode se Se pceceeaced + se a + se
A OES osnetidecaae: rapseeasbancee +
PEVEDTG MACULAL ea taistepnesecsiesa cece: + se + F + + + ae
Wi) IMIR ATO™ Fane scees doecrccec es + Se aha ar <> + ar +
Bi AGAR DIGH Soh aonsosapcaoooneaccandeds ae + +
SFU MLOCULCLLC enna uments cecener sireeiispn + SF + oF
ps) SUDULGLO cc eedorsecacccamcens es + 3B ar + + + +f =e
“4 CHENULOTE nendesiceacax ccceue ence + + + + =n Ts ==
a, CPA Ranh. coatsbanenonsetioontee + ap at
ga) URDYLONGD So woeceen ee se-neeccacne + ae af +. + +
Aa NRLUILULCieamtlesencannedeme ssactlcetns + aF +

MARINE MOLLUSCA,.

AA o8 =| cA Ce)
SES RE) Gea lec em | aa ste ee a
Ss BS ies BO hers | sal se
SPs ao | & 4 Hs
Terebra undulata . + +
oT offinis + + + + + +
fa columellaris. +
Gy RIGO RIG itereoposconoeeprconasts or ar
Fook LQMCNG Grcvauc raven sercyenaneees ot +
Ne) EMD LOLA CONG E unmce annette eecee ct
= LANCENTGE Won .ateaees eae + + 3 a0
~ Célidonvliaiy. ac.
Plewurotoma armillata .........c0cee ee oF
- ULC Reeaeee ceneeonasacecec an
cs CLOTS Rone nen Sacer nee + + of ae + SP + +
IDTLUUG CLASPETALA v.00... -ee een setae ar
Clathurella robillardi 0.0.0.6 cece eee +
Daphnellansatusaua enecuawcenkee skate ap
Oliva erythrostoma ..............0002-+- ze a1 ‘is 3F ar Sr ar a
FLOPPG VENUTICOSG, crccxascnacsesucsen es =e + oF + 36 ae 35 ar ts =F
> MANON esac dscdecssee ce tateearetes + oF + =r Ar i
Marginella picturata v.00... +
WAU EPISCOPAUS ne. encnee cents anes + 3E 32 3F + at +
MMR DOTULLCAIES secomateceet eater ere + + 35 GF
apn CCSBELLAEG, osc dhvencevanvescesoasemee a
Sp) AQUBLD cciccican ese senaorenacnsaecne a at
Fp QUQULGLUES te reacacecmeateee sosseens + 3 TE
Ae MUG. COREL Babe naconceschorrenereccer + + ts ae
Bat eeCLON URS CULM cmerte cate eeeee seer + 35 + cts a5
Ov Wariegatancvs:oe et gions + ar
pn CULUIG RUIN Brace sepocac aera: Sota + P 25 + ata +
G3. A AICI cbopopecberencundassaratocces + ar oo
Boi PUNSCILLDLGueeed. decciecec tetas secon +
aot UUETUN ALG eer aenenarenadeces ane aas: a a7 ate
FET RUCLCTU/2) Meeap eRe ORC ROBEY er ecE DER ce + or + + + oF a ar
Fy PACUMINALA mentasctoudeceescrs+cens + ‘ 1s oF
STE QUT UCLLLOUOLES rantsemean eens snes 35
SSE CLOSDOT CLD tontarancdceaniastestnlaseia ot ota + ch +
55) SCUUPULULS eo ona caatsmrececaceretsne a
Poem LOUDICl im. nometiesesess aes avisiees | mrt + a
See CRE ON Dee ercntoncaseetee tes ecitt
Asie LLCSIQY CS Utirart Becetciead: neceev ore + . or ay
3, angustissima, N. 8p. .........
Cylindra sinensis .......cccee veceeeeeeee a5
Latirus craticulatus.....ccccccccceeseeee + to + + +
By CELI erecaneaceer preceero a oe Fe a5
Peristernia nassatula 00.2.0... + = : + ar + a5 +

76—2

591

592

EDGAR A. SMITH.

a [oe jeg | ee 2 | se lee [aa
= S I 3 | es S 3 oa
BP | 2-2 23 | 3 | eee ee
3 = | B | os z io) S s <5
sen Peal wails ¥ sis
|
Leucozonia smaragdula .............+. aE alla or + + +
Cynodonta cornigerd ......-...22-000+- se ay + + + +
Engina mendicaria ......-..--0.2s00000 + + + + + +
a DONASLD) ve sean nesacaseats cuaawee + + +
+ PAMDSOM im 5. Stes ies tine seals +
IPOS) TOREALUS tes ween neeewseenee= «onsen oe ro . + + ae +
PLEX ERIS Neo teen er ence esa saarie oe +
Niassa btfarid ......2.2.2--.00.cerereses == ap ae
Be RTL T QUUb cs een Wn ecieslemnen= steels + +
Sn IMLOTG Leena te onenenemonda as tesa = + + +
ay GLCHRTERIEE ceSheong) oopccseeasaec0 + + + +
Pen) ECHUNATOe canst nn eoneumnee sees + +
» glans at ue +
» granifera 6 + =
», maldivensis, 0. sp. ..
» mulukuensis, D. sp..... :
aa) CCSULLO “orese Peeeaeceewenwer +
5, subtranslucida, 0. sp..........
ay Ube REE aa¥5 {3} 05 Gopceesceccorce
Columbella conspersa .......0..e0eee ee te
“5 LTS cecebcsoosen eooesan 26 + +
53 Mj OS DUA CO ee crete enter aie 250 +
= WD OLATUTS tree esc eaNocancnae
op CLT CMY ET Umit ear ect eiene enc + + 35 +t + + +
AG MOPUONS cana otsanacansiess + + ae
Murex ternispina .......-.....0.+-- +0002 ar 35 + +
“5 TEDNISTEUMI Bee teense aa ceeiee « Sr + 3° + +
x UPUPLCT;UB) canoes danas codecs er 35 2 + 3
a CULEMLUS, cohen cceeeh sasher esos as +
An COUSTUBS dxtae de cotta etenoaanientaee ar + - + + + =P
a TUOLGUILOSUS saeceucreexaseeseeeee + 53
3 TAMOSUS 2.02.2 c00csreerereverecrens ay + ot at = ap +
Hi SQUMONEUS 500-5 on on naenncanenenrs 5 .
=F pleurotomotdes .........-+.-+0++- .
a SUDMISSUS, TNS HPs..vbssesess ee. 7 ra
PUrpuUrd APMIGETA.. 622.602 cecerveereeeees + + = = ae a5
PA bitubercularis .........--++-- te
.3 NippocastanumM «2.0.6.0. + ar a5 ts nF ae 35 SF
50 TOG, © cecgcbi IIIT + a5 + ves +
TOPA8) SUHULG) v.22 .cecaceosctennoneennenaa= ate FF
Sa ROTDUNE coca postman tees sieaaies edn = + 35 an =: =f +
Sistrum digitatwm ..........0.eseeeees + + af “2 +
a ROY COUT. option vevsomen annie + + + a9 Se iets + +

MARINE MOLLUSCA.

x se) 8 a 2.8
2/2)2/42\22)/8/ 2/2 | |s8
SP ee ieee Weel Piles (A te Ih tents |e eine eee eared
BO || te FI Ea || ice |] Set |) teh |) a |<
Sth cabal adh paces w Hes ns Ep) Gamat
mo | A =
SUSUMU MUCINUS: caves | ecnwanwece<deeelene + + + a5 ar an a5 AF
my tuberculatum ......0..c.ece00s + + + + + + af ats
A UNGGCUNY .: tga seaseeeeeroeate ssn + > + + an au Wi
+ CLACMIN:..cc.ssacpetaa sess aceite tes a0 45
ry SPUMOBWIH 00 ccuscervesneuson-ss eee a
= DicOntCUinicceerose cece eoae : ste ay
a CAVETNOSUM....-.00c0e0es2e0eee a
8 LOBLOMO ab spanaceneesetaat Res: > 7 “
in SQUCTOSIM epee saestenes a
A concatenatum + ar ae
As decussatum..... +
Coralliophila monodonta.. + +f et
A neritoidea . + + + ar oF a5 ar
4 SUtUT AUS eee eee ok a
as squamosissima ..
Leptoconchus ellipticus ....... .. +
0 CUMANG Wane ease fo
Lotorium chlorostoma ...............++- + + + ar : a a
es GEMMALUM .....2--2eeneen ses + + ar ate
be FAOPOUIE: conceceBounseseconcebene + ‘ +
a CUDEOSUM ......ceececeeeeeeess + + + ar ar ar t ar
Colubraria ceylonensis..........00.200+6 : ar oF at
es ODSEUT OD eatnesueneveetmerees + : ar oe
“; DTACtCQLa, yeccnsteccaere-ees a : + + ar
i Gigitalem@arceste te aeceoe oe . + ats
Distortrid QNUS....00...0c0cereeeeeesenens + + é + + ae at
Ranella bufonia ...........000+ acces + + # + a 20
Na) AGTOMISENGs <2, coavnisszonensce + + 35 oF + at
rp LOM US eeaweeseeece ees near tere + + + + + sr 7 +
3 tuberosissima .........00ee00es + aur
iy EROMGE creme ade easeeeee east + a . os .
CGSStGACONMULR Meee ase assetesear ees + ate =e
Pe MUG eee ree aee nencestee + + + + + fe oF
Malea pomum ........c0eeccceenceseeeier + + ar ate ar
Doliwm costatunm ....:0-<0c0.c22cse02eees + - =F
5 OUTRUN: Gradessnsebece aCAnconeen, + + + + + + ar
Pirrila, ficotdes .......-.con.ce-sseeererees + + a ar see
Cypraa annulus .........0cccceeeseeeees ~ + + 3 3 + + aF cfs
nn AT UOUC EE coor ee era + + + a + + ar as ate
5 CORSO co picdeicn Se PPORGELEC OEE + - + + ae + ap
és QEELULS Wet ene eee + + ae + ae ae ar +
- caput-serpentis ...........666 + + + ot =r t- + ot;
a CONTIEOLON wee scvscaeseeresgrm=-5s + + se a ae + aF ae + a5

593

594

EDGAR A. SMITH.
a | 8 |e \te |S) 2 eee oe
Be} os | | at) ol Seal oem ene teetaes
5 a =) 3 |nsd!] @ > q = aS
a ao} 8 Bl gh a to) = Se) Stet
SA Suis Ae alse ce alee er
(GN OHAT CURUTTGH, “Ceeseonesee-ce- ceded + + Ste ae ae + + oh ah
es (HIT OS ESOT ee ecceonAcacascatboe =F a> ele
a CTUDT ATE. somes stoes cece eens eas + ieiate at aT
56 CTOSG: ie. sce wenees vice cevoes ere 3 + Se + oF + + ++ ate sr
oe |) iLLLOS Cpa ne: eeRCEEe + a a a5 et
FAM IUENGTENOS ee ce ns see ee an + ny
rs GLOBULES cent tern case ee cet - ae a a
a ELVOUMD ninmema sade aetineeeere + + a= + + + + + ae
a RUUD aa te ts heaps ate + ar or +
A ER LOGLLC ecto Gets gent aa ae + + + + + ate as + as
A (TITTIES Sten Gepcocutorodse se + ie
a YING Meee osname eke acre + + + ct + + = + Fa
os MOU ULONGIG. eee sa aeaeenee ks + =P aF + + + + a7
5 IUNOTULONG teach ee bance ee Sr of a
AA MROTLEL Le steerer ns ae selena es eaiced oak se + + + + + se a5
* TORTUN SID © Ase nBoaceenre RE Rbreme + “h ap ais
4 BEODNY MBO mo eeeiancteeceee acne + + + + + + ae
A CAUD Oar ere ese op ee + + =F + + + ar ar
a At fk PE AOE SPC ent Ree AB + + + + + + ts a
9 WOULELUUS meters cncnde tases se eeae aR + ar + GF + ae aF ai
_ RUDUKET Unt Pe ae eee oe 7F F
an PACE Oba cod eats ae ot aeons + = + ate
DPrivia pelluctdula ........cscssseesees- a +
Strombus dentatus .........60..00c00 00 ae =F “
+3 WflOridus -s......:. + + + + + ats
a gibberulus ..... an + + 3+ + 55 + ar + at
labiosus...... F a
oF lentiginosus . 45 st + SF +
a (RTUATRIDES Veapeone tooo ee apere ae ot
3 [eC COR Sbapbaceeeer te anee sh: a5 +
TOG IRED [ETO secmne nemeepcendbecsaone a + ae ae ae + we
7 COU IRATIROS Soe cacecendecyetinson + + + + aF + 3F
Terebellum terebellum ..............0665 + +h ar =F + tee
SPraforis, COTMGATUAY -easceesneenessseieee ay ag
4 CLEGUNS or. ds fer contends Saten seas +
Fane ASCII PULLS Bek romeane doce sees Ree 2 ae oo
sy lilaceocinctus, 0. Sp. ...... + oT
y CL CELLS at AD a Beeecen eee nes
a QTACULE, Del SPsp--tecreseeese ss
oa) | ROUTE, MDS SD slr acaecgsaaneasees 20 09 .
Vertagus articulatus ..........c0.00005 n60 + oe
Fri TROON) eesice-eemeniocence san 7 + + + ae se Sta oe
rh KOCK ae a Misncevated eae Cees + + a5 ac

MARINE MOLLUSCA.

lee | see lee les re ee
ray Weg, leceh Wea |e OS CS. oe ee ee
Be Weep 6 Be rss Wears aes ik eee en hs alee
G se | S | os 2 12) =} S | <a
Brat Sl Vee ae |e Se Were
Vertagus obeliscus = f 3 ar + + + +
Cerithium colwmna + + + +
f ECTLUNAUCUM Me wtecenecatentene 3F + + + +
mo LACTCUTIS one neem ecm eiene ssed| a2 ar +
a PUPCTUUM, svar ccneneeesserens “+
ie CALEB OSIM: Jocces cen eeennens De is
x tuberculatum ......c.0.c00 + + +
a ALDOVATICOSUM,..002. 0 secs ct
Terebralia palustris ...........c0eceenens + + oF + + +
VO OAUIUS, COGUUT Cc cen ceree cca eoes- <2 as =F + se ae + ae +
IPLIOMOGZIS ANEDCUS: w.hacenstescespnenessa +
Littorina glabrata .......0.eeecee eee ee + + + + + +
a4 BCADTO! ease tance deere oo + + + + + + ar
os Und nlata onscreen + + he
Solarium modestum ... + ar
Philippia cingulum ... + + + a5
Mitrularia cicatricosa .. + +
tr CQUESUIIRN, eveee see eteaseden + + Te +
Hipponyx danieli............0...c0ess00 +
a QUSUF AUS arse casece ooeeaaes + + + +
Thyca crystallind.........c.cervecseeeees ar +
KXenophora iNdicd.........-.secereveeeees + +
Vanicoro cancellata .........c.cc0ceeeees + + + a
Natica robillardt .........ccceeceeseeeees +
ag = EDEL? epprccococnsonsneceeGnnoccd + ae
Polinices, MAMIUlA....--c2.2s-<¢--<-enseee + + oF + + + + ar + =
“5 QUDUMET cv hperecase en sensntes + : to
:; MELANOSTOMA...... ee eeeeee vee + + ar 35 fe + + 3F +
ie SUMIGE » veces antag nee ncuneemese + nn +F 5° ae ae ce
Tanthina communis ..........0.00.00008 = + + ae nF $F oF —
Scala replicata........ 2 + +
Ben COMROTS = cece +
>, texta, N. Sp.
Pyramidella corrugata ........... occ! SF
eA nodicineta ........+ 2eceo4 oe ae oe
Obeliscus dolabratus............---.-++++ + + + + + + =
“Sy MOTIIIS: eae ttaschcet- aca 5tks +
a SULCOTUB Sc feccasicensoruceos tees + + + + + + +
Neritd QUbtC ld s....<2..--acccnssoecoveeee + + ef + + ss =f + +
PAU HUSURLOmrranteneeaneeecadare ese es a +
AMR DLC OLE ree cnet ean saeaectee tues + - + + + + +
eo PM POUL ace cer een - + + - ~ + - + -
Turbo ArgyVOstOMa ....2..00-020eeeeee +5 + + a no

595

EDGAR A. SMITH.
ra aa — a 23
bee bebeibe so) ee
a Do 2 q Se EI = g a ene
he merubcbes meeps |e ls las
eet Pee: a |" las
Liotia squamicostata, 0, sp..........
Trochus UMcrassatus..........000c0s00+0 sf ae
a WIUGCULCLUSS eons density asisrentees ate A + a
7 QUESUB i os s-<cmesueeene
A radiatus ~ + + - a
Thalotia maldivensis, n. sp..........
Clanculus Atropurpureus ......00.00000: ar at
Perrvinta, GNGWASET .cb.ccceeeeasesseees +
Forskalia pulcherrima ...........02.04.+ +e 35
Monilia calycutus........0..c1..e6ecereee + or
ay BUITLULAIUS eocein Mlecseechesee eee Te
Ethalia rhodomphala, n. sp..........
(CUE) CTO TTL nomeedaceond ac HeoNebeI0907 as
PLQUOTISNOUIN Gece scan ces sen cetr sce eer a oo
Glyphis bombayand .........ccceeeeeeees
aS SINGAPOVENSIS ... 0... cececeev eee +
Emarginula (ncisura .......6..0. 0.000
* POLQTULOC weccee tem cceiee sip
ORO MILES oes vacteasee wen decane seas ea
Schizochiton incisus.............. +
Callochiton platessa........ . ats
Ischnochiton hululensis, n. sp. ...... 5c
a feliduensis, Nn. sp.......
a maldivensis, N. sp. ...
Acanthochites laqueatus ............... aF at
Cryptoplax DUTVOWE .........c0ec eevee eee ar
Proneomenia australis ...........000.00 20 2s
Spondylus imperialis .........0.0.0000- ats
Lima fragilis a Br
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APEC UULLM OBC bess te ceuraclosisiesaesnielceen + + its ae
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a UAT NORTE or ecopbasetesocdr ane
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Semipecten forbesianus ......-....... sat et

MARINE MOLLUSCA.

H ga = a 33
flere aay eesioad| See al cosa eal) ork eta) acon ries
e s g B | oe a oD = s 4a
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a IAURCE pokes AA cE
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IPUNTGNMUIUCAEA ~. nccuecsceeeseseceuine
AEE TUONO s das ccae uatoceniec arene + + + a0
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i QUMICULALA cone net ee + aby ata a0
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as MALACCANUS .......02 00.00 ar
an MUASUUUS ns nwescnandtestccaser
“A UCL ec BRACE BE PEGE Chace + ae ar a
Modiolaria argented ............00.005
Septifer bilocularis .............2.000++- + + ar an IF ap
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pap MAOMIUNGENSIS,. eewatcretvaresieecncee ata 37
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Bs LLEMELLGrnihneesorcsaecsgnedirorecnes : a0
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Thyreopsis coralliophila ............... + xe
Tridacna cumingii ...... dapereccacened a0
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ce MECULOSUM Ns ca. scece seccoacsees . “
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kis levisulcatum, 0. sp. nS cee
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BS DUBIN GLE coccensveoventvetservaxe + + + + +
OUARAIUM iic...ccecascseces fs an
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IL.

77

597

598

EDGAR A. SMITH.

Cardium fornicatum

GRAM GURESU (yc dacccvuxcensosceencccses
= RIRUVICO UE raineht ees cevaecrectens
Meiocardia lamarckii
Trapezium oblongum
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as GLAU ALO) Neccsssncsscnsessdeseee ss
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+
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MARINE MOLLUSGA. 599

It has been noticed in other groups of animals that the specimens collected at the
Maldives have a tendency to be smaller than those found upon the shores of the mainland
and Ceylon. This peculiarity also to a certain extent is noticeable among the Mollusca, or,
at all events, the representatives of many species are small in comparison with the larger
dimensions they often attain.

No attempt has been made to give full synonymy or many references to illustrations,
but it has been thought advisable, in each case, to quote a figure or a description, so as
to show what species was really referred to. This is necessary in some instances where an
author's species has been differently interpreted by his successors, as, for example, in the
case of Strombus auris-dianae of Linné, where Reeve’s and Deshayes’ view of this species
is quite the reverse of that held by Sowerby.

In the account of the numerous species hereafter mentioned the localities where they
were obtained are referred to under numbers only which are explained in the following list
of stations.

I. Addu Atoll, 5—40 fathoms
II. Suvadiva Atoll, 5—44 3
Ill. Haddumati Atoll, 38—40 $
IV. Kolumadulu Atoll, 1—40 a
Ve S. Nilandu Atoll, 1— 36 z

VI. Mulaku Atoll, 1—40 é.
VII. Felidu Atoll, 1—35 5;
VIII. S. Male Atoll, 30) a

IX. MHulule Island, 1—6 es

X. N. Male Atoll, 1—35 7

XI. Goidu Island, On reef

XII. Mahlos Atoll, 4—24, Bs
XII. Fadifolu Atoll, 2—23 :
XIV. Miladumadulu Atoll, 3—28

XV. Mainikoi, On reef and 5—7 fathoms.

[The editor regrets that the dimensions of the whole work on the Fauna of the Maldives
and Laccadives have made it necessary to follow the method adopted by many publications
in placing the list of literature at the end with reference numbers to the same in heavy
black type, and more especially since Mr Smith had most carefully prepared his manuscript
with the references in full under each species. He also deeply regrets that the special
geographical distribution given, in the manuscript, with each form, and which entailed
considerable research into conchological literature, has, for the same reason, had to be omitted.
Roman numerals in italics refer to the volume of the work quoted. Ep.]

cc

600 EDGAR A. SMITH.

Family ACTAEONIDAE.

1. Solidula glabra, Reeve, var. 37, XV. (Solidula sulcata, Pilsbry (nec Voluta sulcata, Gmelin),
34, XV.) X, XV.

In my opinion this is not the Voluta sulcata of Gmelin, as stated by Martens and Pilsbry.
The name su/cata practically proves this, and a reference to the grooved figures in the Conchylien-
Cabinet upon which the species was founded also shows that it is quite distinct from Reeve’s
Tornatella glabra. Moreover Martini (Conch. Cab. vol. 11. p. 124) described his shell as being
“tief gefurchet,” a character not at all applicable to the present species but admirably suiting the
well-known Solidula solidula (Linn.) with which, in my opinion, the V. sulcata is synonymous.

The specimens from the aboye stations are small and slender.
2. Solidula nitidula, Lamk. 34, YV. IX, XV.
3. Solidula solidula, Linn., var. 34, XV. VIII.

A single specimen, pure white, with only a few grey dots upon the ridges near the outer lip.

4. Solidula tessellata, Reeve, 34, XV. IX.

Family BULLIDAE.
5. Bulla ampulla, Linn. 34, XV. II, IX, XI.

Family AKERIDAE.
6. Wolvatella cincta, Nevill, 34, XV. XV.

Family SCAPHANDRIDAE.
7. Atys cylindrica, Helbling, 34, XV. II.
8. Atys naucum, Linn., 34, XV. II, III, V, IX.
9. Atys succisa, Ehrenberg, 40,a, XV/I.,1,c. XV.

It is uncertain whether the shells figured by Adams and Sowerby truly represent Ehrenberg’s
species, but it is equally certain that they do not belong to A. cylindrica as stated by Pilsbry.
The latter has made a curious mistake in connection with A. swecisa (Man. Conch. vol. xv. p. 267).
He states, “There is the trace of a median gibbosity,” whereas Ehrenberg distinctly says it is
absent, “gibberis vestigium in zona media deest.”

Family OX YNOEIDAE.
10. Oxynoe delicatula, Nevill, 34, XV. XV.

Family PHILINIDAE. .
11, Cryptophthalmus minikoiensis, n. sp. (Pl. XXXV. figs. 1, 2). XV.

Animal corpore oblongo, angusto; discus capitalis parte corporis reliqua brevior, bipartitus, parte
anteriori scutiformi, antice bimarginata, truncata, postice angustata, bilobata, parte posteriori postice

MARINE MOLLUSGA. 601

truncata, ad latera carinata et lamellata; pes infra orem leviter sinuatus, ad angulos rotundatus,
extensionibus lateralibus semicircularibus, sed in exemplo singulo supra tergum reflexis ; testa minima,
ad extremitatem corporis posticam posita, cute tenuissima tecta, tenuis, semipellucida, supra convexa,
semiconvoluta, postice acuminata, striis incrementi tenuibus sculpta; branchiae plumiformes, multi-
Jamellatae, sensim versus extremitatem attenuatae, infra testam sitae. Longit. tota 17 wmillim.
Discus capitalis 7 longus, 4 latus. Diam. lobis pedis expansis 12 millim. Testa 4 longa, 21 lata,

Closely allied to C. smaragdinus, Leuckart, and differing only in the form of the shell. This
is exactly the same as that erroneously figured by H. and A. Adams in the Genera of Recent
Mollusca, vol. 11. Pl. tvin, figs. 3a, 36, as Chelidonura hirundinina of Quoy and Gaimard. A
comparison of these figures with those of Leuckart' and Ehrenberg? at once shows the considerable
difference in form.

The shell of Crypt. Juteus, Quoy and Gaimard, is more like that of the present species, but
the short folds on each side of the head-shield do not appear to be present in that species, at
all events they are not referred to by the authors. The most remarkable character in this species
consists in the structure of the cephalic disc. It is, as it were, in two parts, the one superimposed
upon the other. The anterior portion is shield-shaped, being broad, truncate, and double-edged in
front, and narrowed behind. The converging sides are not, however, united posteriorly, but form
very small lobes, attached to the hinder portion of the disc. The latter is squarely truncate behind,
keeled at the sides, and exhibits beneath the carinate margins about fifteen short oblique lamellae.
This is a very striking feature, and occurring in C. smaragdinus, makes one doubtful whether
the present species, although the shell is very different, is really specitically distinct.

Family CONIDAE.
12. Conus arenatus, Hwass. 37,7. VIII, IX, XI, XII, XIV, XV.
13. Conus betulinus, Linn. 37, 7. (C. medusa, Gmelin, 16) IX, XI.

14. Conus catus, Hwass. 37, J. (var.=C. discrepans? Sowerby, 40, a, var. = CU. adansoni,
Reeve (now Lamk.), 37, /., var.=C. nigropunctatus, Sowerby, 40,0, ///.) IX, XV.

15. Conus ceylonensis, Hwass. 37, J. (var. =C. pusillus, Chemnitz, 37, Z., var.=C. nanus,
Broderip, 37, 7.) IX.

16. Conus distans, Hwass. 37,7. (C. mennonitarwm coronatus, Chemnitz, 9) IX.
17. Conus eburneus, Hwass. 37,7. IX, XII, XV.

18. Conus ermineus, Born. 40, /, ///. (C. lithoglyphus, Meuschen, 37, J. C. orleanus, Bolten,
30, var.=C. lacinulatus, Kiener, 23, var.=C. carpenteri, Crosse, 11, a) IX.

19. Conus flavidus, Lamarck. 37,/. (C. maltzanianus, Weinkaufl, 43, var. granulata=C.
frigidus, Reeve, 37, 1., var.=C. neglectus, Pease, 31,5) XV.

20. Gonus generalis, Linn. 37, J. (var.=C. maldivus, Hwass. 37, [., var.=C. monile, Hwass.
37, 1, var.=C. bayani, Jousseaume, 22, jun.=C. spiroglowus, Deshayes, 40,6, X//.) XIV.

21. Conus glans, Hwass. 37, J. (var.=C. tenwistriatus, Sowerby, 40,0, ///., var.=C. fabula,
‘Sowerby, 37, Z., var.=C. scabriusculus, Chemnitz, 9) LX.

22. Gonus hebraeus, Linn. 37, 7. (var.=C. vermiculatus, Lamarck ; var. =C. chaldaeus, Bolten,
30) IX.

1 See Pilsbry, Man. Conch. vol. xvi. Pl. vi. figs. 33-34. * See Pilsbry, lc. fig. 32.

602 EDGAR A. SMITH.

23. Conus lividus, Hwass. 37, /. (var.=C. citrinus, Gmelin, 40, b, ///., var.=C. sanguino-
lentus, Quoy and Gaimard, 35, var.=C. mawrus, Gray? 18,5) IX, XI.
24. Conus miles, Linn. 37,/. IX.

25. Conus minimus, Linn. 37,/. (Conus coronatus, Dillwyn (non Gmelin), 43. C. barba-
densis, Hwass. 40,6, /1I. C. aristophanes (Duclos?), 40, 6,71.) I, XV.

26. Conus nussatella, Linn. 37, /. (C. terebra, Chemnitz (nec Born), 9). IX.

27. Conus pennaceus, Born, 37,7. IX, XI.

C. praelatus, Hwass, C. rubiginosus, Hwass, C. episcopus, Hwass, C. omaria, Hwass, C. magni-
ficus, Reeve, (. elisae, Kiener, C. racemosuws, Sowerby, and C. madagascariensis, Sowerby, may be
regarded as varieties of this species.

28. Conus quercinus, Linn. 37,/. (C. buaeus, Link, 30. C. ponderosus, Beck, 6). II.

29. Conus consors, var. 40,6, ///. (var.=C. innexus, A. Adams, I. 6) I.

With this species, besides C. innewus, I also associate, as varieties, C. anceps, A. Adams and
C. daullei, Crosse.

The single specimen from Addu atoll differs from the typical form of the variety imnexus in
being almost entirely white, and of a slightly more slender form. It is clothed with a thinnish
greyish periostracum and the upper whorls of the spire are finely coronated and of a lilac tint.

30. Conus tendineus, Hwass. 40,7. IX.
31. Conus tessellatus, Born. 37,7. IV, V, IX, XIV.
32. Conus textile, Linn. 37,/. IX.

The variation in this species is most remarkable, and although we can recognise typical examples
of the many named forms, personally I have not been able to accept as distinct species the
following :—C. vicariws, Lamarck, C. verriculum, Reeve, O. textilina, Kiener, C. tigrinus, Sowerby,
C. corbula, Sowerby, C. scriptus, Sowerby, C. panniculus, Lamarck, C. canonicus, Hwass, C. rubescens,
Bonnet, C. legatus, Lamarck, C. paulucciae, Sowerby, C. archiepiscopus, Hwass, and C. abbas, Hwass.

33. Conus tulipa, Linn. 37,7. (var.=C. obscurus, Humphreys, 37, 7, jun.=C. borbonicus,
Adams, 2, 42). IX, XI, XV.

34. Conus zonatus, Hwass. 37,/. IX.

Family TEREBRIDAE.
35. Terebra maculata, Linn. 37, X//. IX, XI, XV.
36. Terebra argus, Hinds, 37, X//. (Terebra nebulosa, Kiener, 23). IX.
37. ‘Terebra dimidiata, Linn. 37, XY//. IX.
38. Terebra oculata, Lamarck, 37, X//7. IX.
39. Terebra subulata, Linn. 37, X//. XI.
40. Terebra crenulata, Linn. 37, X//7. XV.

Buecinum varicosum, Gmelin, B. luteolum, Martyn, are synonyms and 7. jfimbriata, Deshayes,
and 7. interlineata, Deshayes, I regard as varieties of this species.

MARINE MOLLUSCA. 603

41. Terebra cingulifera, Lamarck, 37, X//. (Buccinum suceinctwm, Gmelin, 16. 7. punctu-
lata, Sowerby, 40,c. TZ. punctatostriata, Gray, 18,0. TZ. pallida, Deshayes, 12,6. T. chinensis,
Deshayes, 12,d). IX, X, XII.

42. Terebra babylonia, Lamarck, 37, X//. XIV.
43. Trerebra monile, Quoy and Gaimard, 37, X//._ II.

44, Terebra (Myurella) undulata, Gray, var. 37, X/J. (7. approximata, Deshayes, 12, ¢).
1) Sa

45. Terebra (Myurella) affinis, Gray, 37, X/J. (7. striata, Quoy and Gaimard, 34). IX,
EXC exe OXCLP TS XOV),

46. Terebra (Myurella) columellaris, Hinds, 37, X//. VIII, X, XII.
47. Terebra (Myurella) bruguieri, Deshayes, 37, X/J. (7. hindsii, Deshayes, 12,6). II.

48. Terebra (Myurella) amoena, Deshayes, 37, XJJ. (7. andamanica, Melvill and Sykes,
39). VII.

A comparison of the types of the above-named species shows that they are merely different
stages of growth of one and the same form,

49. Terebra (Myurella) violascens, Hinds, var. 20,), 37, X//. II, IV, VI.

The specimens from the above stations are not typical and vary inter se. None are of a violet
tint, but whitish or yellowish. The shell described by Reeve had the whorls undivided by a groove
or punctured depression, whereas Hinds described the type with whorls ‘“superné lined impressa
obsolete cinctis.” The Maldive specimens agree with the type in this respect. Terebra exigua,
Deshayes, and 7’. polygyrata, Deshayes, are probably only slight varieties.

50. Terebra (Hastula) lanceata, Linn. 37, X/Z. (var. penicillata, Hinds, 37, XJ/J, var.
venosa, Hinds, 37, XZ/.). XV.

The specimen from Minikoi (station XV.) belongs to the variety penicillata.

51. Terebra celidonota, Melvill and Sykes, 29. XIV.

Family PLEUROTOMIDAE.
52. Pleurotoma armillata, Reeve, 37,7. VI.
53. Pleurotoma bijubata, Reeve, 37,7. VIII.
54. Pleurotoma tigrina, Lamarck, 37,7. VI, IX, XIV.
55. Drillia (Clavus) exasperata, Reeve (Plewrotoma exasperata, Reeve, 37, Z.). II.
56. Clathurella robillardi, Adams, 41,VZ. XIV.
57. Daphnella saturata, Reeve, 37,7. II, VI.

Family OLIVIDAE.
58. Oliva erythrostoma, Lamarck (var. ponderosa, Reeve, 37, VZ.). IV, VI, IX, XI.

Apparently browsing on a black crinoid (Gardiner).

604 EDGAR A. SMITH.

Family HARPIDAE.
59. Harpa ventricosa, Linn. 37,/. IX.

60. Harpa minor, Lamarck, 37,/. IX.

A specimen captured by Mr Gardiner at station I. has the hinder portion of the foot detached.
Judging from the large size of the separated portion, the animal apparently divested itself of it,
being unable to withdraw this bulky appendage within the shell. Or, may it not be cast off as
a peace offering to any fish or other predatory creature who attacks the helpless Mollusc?

Family MARGINELLIDAE.

61. Marginella picturata, Nevill, var. Pl. XXXV. figs. 3, 4 (JZ. picturata, Weinkauff, 43). X.

The two specimens which I believe to be a variety of this species differ from the type in
having the outer lip strongly denticulate within, in one example the denticles numbering thirteen,
in the other only nine. Both are of a fleshy tint, with two white narrow zones with reddish-
brown spots upon the penultimate whorl, and three upon the last, one at the suture, one at the
shoulder, and one towards the base, all terminating upon the thickened labrum in a red _ spot.
Between the second and third zones there are three or four transverse lines of pinkish red colour
and the extreme tip of the spire is also reddish. The brown spots upon the narrow white zones
are small and might be described as cross lines rather than spots.

Family MITRIDAE,
62. Mitra episcopalis, Linn. 40,),/V. XI.
63. Mitra pontificalis, Lamarck, 40,0,/V. IX.
64, Mitra tessellata, Martyn, 40,6,/V. IX.
65. Mitra adusta, Lamarck, var. 40,0,/V. IX.

One specimen of a uniform rich brown colour and without any crenulations beneath the sutures.
66. Mitra digitalis (Chemnitz), Dillwyn, 40,6,/V. IX.

67. Mitra (Scabricola) crenifera, Lamarck, 40,6,/V. IV.

68. Mitra (Scabricola) scabriuscula, Linn. 40,),/V. XI.

69. Mitra (Scabricola) variegata, Reeve, 40,6,/V. II.

70. Mitra (Chrysame) cucumerina, Lamarck, 40,0,7V. IX, XIII, XV.
71. Mitra (Cancilla) filaris, Linn. 41,//. (J/. filosa, Born, 40,6, /7V.) XI.
72. Mitra (Cancilla) insculpta, A. Adams, var. 40,),/V. V, VI.

73. Mitra (Cancilla) interlirata, Reeve, 40,},/V. II.

74. Mitra (Strigatella) literata, Lamarck, 40,0,/V. IX, XI, XV.

75. Mitra (Strigatella) acuminata, Swainson, 40,),//. I, IX, XI.

76. Mitra (Strigatella) auriculoides, Reeve, var. 40,0,/V. IX.

The two specimens obtained by Mr Gardiner differ considerably from all the figures given of
this species. They are much more slender, and the upper part of the shell above the white zone

MARINE MOLLUSCA. 605

is white, blotched with dark chestnut. The lower part of the body-whorl is minutely white-dotted.
Intermediate forms connecting this with the type occur in the British Museum collection. Length,
17 mm., diam. 7 mm.

77. Mitra (Costellaria) exasperata, Gmelin, 40, }, JV. (var. =. arenosa, Lamarck, 40, b, [V.)
AVéliemVeleeIEXe, OX] XEN: SXDV, SV.

The two specimens from station XIV. are variations of the form arenosa, differing from normal
examples in being almost entirely white, with only one or two faint zones below the middle of

the body-whorl.
78. Mitra (Costellaria) sculptilis, Reeve, 37,//. II, V, X.
79. Mitra (Costellaria) acupicta, Reeve, 40,0,/V. V.

80. Mitra (Costellaria) armigera, Reeve, var. 37,//. IX.

A single specimen only of this species was obtained. It differs from the type in being whiter,
with a thread-like brown line round the middle of the upper whorls and three on the last. The
locality of the type is unknown. J. turrigera, Reeve, is closely allied, but more sculptured
between the costae.

81. Mitra (Costellaria) deshayesii, Reeve, 40,4,/V. X.

82. Mitra (Costellaria) angustissima, n. sp. (Pl. XXXYV. fig. 5). V.

Testa gracillima, elongato-fusiformis, pallide fuscescens, ad partem anfractuum superiorem albida,
fusco maculata, et circa medium anfr. ultimi maculis subquadratis notata; anfractus circiter 10,
superiores subgradati, fere plani, caeteri leviter convexiusculi, costis numerosis (circiter 28) gracilibus
instructi, et sulcis spiralibus inter costas punctatis (in anfr. penult. 7, in ultimo 12—13) sculpti,
ultimus antice cirea caudam sulcis obliquis fortioribus haud punctatis ornatus; apertura parva, angusta ;

longit. totius 1 paulo superans; columella plicis.quinque obliquis instructa, callo tenui induta; Jabrum

tenue. Longit. 12}mm., diam, 4. Apertura 5 longa, 14 lata.

Remarkable for its very slender form and the punctate striae between the costellae. The folds
on the columella correspond to the oblique ridges between the sulci on the cauda of the body-
whorl. These ridges are spotted with pale brown.

83. Cylindra sinensis, Reeve, 40,),/V. VI.

Family FASCIOLARIIDAE.
84. Latirus craticulatus, Linn. (Zwrbinella craticulata, Reeve, 37,IV.) IX.
85. Latirus fastigium, Reeve (Zurbinclla fastigium, Reeve, 37,/V.) XII.
86. Peristernia nassatula, Lamarck (Z7wrbinella nassatula, Reeve, 37,/V.) IX.

87. Leucozonia smaragdula, Linn. (Zurbinella smaragdula, Reeve, 37,7V.) IX, XI, XIII.

Family TURBINELLIDAE.

88. Cynodonta cornigera, Lamarck (Vurbinella cornigera, Reeve, 37, ZV.) Maldives.
G. IL. 78

606 EDGAR A. SMITH.

Family BUCCINIDAE.
89. Engina mendicaria, Lamarck (/icinula mendicaria, Reeve, 37, /1/.) XI.
90. Engina bonasia, Martens (Plicatella (Peristernia) bonasia, Martens, 26). IX.

91. Engina rawsoni, Melvill (Sistrum rawsoni, Melvill, 27,a). V.

The columella is described as ‘“‘simplice,” but all the specimens examined, including the type,
exhibit a thin circumscribed callosity, with a small transverse tubercle at the upper end, and a
few anteriorly. All also have a pale peripheral zone, and some of the costae are whitish. In
one of the Maldive specimens, white is the predominating tint; on the contrary, a second example
is almost entirely of a rich brown colour, excepting the whitish zone round the middle of the
body-whorl, which is rather more contracted and produced anteriorly than usual, the canal, in
consequence, appearing a little longer.

92. Phos roseatus, Hinds, 40,3, ///. II, IV, V, VI, VII.
93. Phos textilis, A. Adams, 40,,//Z. V, XIII.

The specimens from the above localities are considerably smaller than the type from the
Philippine Islands but do not offer any tangible differences in other respects. The protoconch is
large and consists of three smooth glossy convex yellowish whorls. The normal volutions may vary
from four-and-a-half to five-and-a-half.

Family NASSIDAE.

94. Wassa bifaria, Baird, var. 5,39, 7 II, IV, V, VII, X.

The Maldive specimens belong to the same variety as those from the Andaman Islands which
I recorded and figured at the above reference. Tryon in his absurd craze for “lumping” considered
this species “a stumpy variety” of .V. hirta, Kiener, a species with which it has in faet but
slight resemblance, differing in size, colour and sculpture.

95. INassa marratii, Smith, 39,¢,f XI.

Two specimens were obtained agreeing with the type excepting in their larger size, being
19 mm. in length, like the Andaman examples figured in the P. ZS.

96. INassa monile, Kiener, 37,V///. XI.

97. INassa stigmaria, A. Adams, 37,V///. V, XII, XV.

98. Wassa echinata, A. Adams, 37,VJ///. IX, X, XII, XV.

99. INassa glans, Linn. 37,V///._ III.

100. Nassa granifera, Kiener, 37,V///. IV, VIII, X, XIII, XV.

101. Nassa maldivensis, n. sp. (Pl. XX XV. figs. 6—8). VIII, IX, XII.

Testa WV. alyidae’ similis, sed concolor, albida, anfractu ultimo majori; lirae aperturae circiter
12 et columellae rugae validae. Longit. 25mm., diam. 15. Apertura cum labro 15 longa, 7 lata.
5 fo} ? fn)

var. a. VII. Testa minor, anfractu ultimo costato, Longit. 20 mm., diam. 11.

Shell ovate, turreted, white; whorls 64 gradate, upper ones costate, the costae nodose at the
upper end through being traversed by a depression or groove, gradually dying out so that the

1 Nassa algida, Reeve, Con. Icon, vol. vit. figs. 1454, b.

MARINE MOLLUSCA. 607

body-whorl is quite smooth excepting towards the labrum where a few sulci parallel with it are
sometimes visible and four or five rather coarse grooves encircle the base. The outer lip is sharp
at the edge with four or five denticles anteriorly thickened and lirate within. lLirae twelve in
number, produced within the aperture, thread-like, not reaching to the edge of the lip. Columella
transversely lirate, furnished with a free-edged callosity. The operculum is yellowish, unguiform,
with a terminal nucleus, and serrated on each side towards the apex.

102. Nassa mulukuensis, n. sp. (Pl. XXXYV. figs. 9, 10). VI.

Testa WV. maldivensi similis, sed gracilior, anfractibus superioribus spiraliter liratis, operculo
diverso instructa.

Longit. 24 mm., diam. 12. Apertura 12} longa, 6 lata.

Shell elongate-ovate, turreted, whitish, with a few dots and spots of a reddish colour on the
upper edge of the whorls, especially on the dorsal margin of the body-whorl. Whorls 9, the three
apical smooth, glassy, the four following obliquely costate and transversely sulcate, the grooves
causing the costae to be somewhat nodulous. The last two whorls are smooth, excepting the
subsutural row of nodules and a few sulci parallel with the labrum and around the lower part
of the body-whorl. The labrum, lirae within and the columellar callosity are as in WV. maldivensis.

The operculum, however, is quite different from that of the species referred to, being ovate,
concentric, with a central nucleus, and having only a few serrations on one (the outer) side.

103. INassa subtranslucida, n. sp. (Pl. XX XV. fig. 11). V.

Testa elongata, turrita, subpellucida, nitens, albida, maculis et lineis dilute fuscis irregulariter
picta; anfractus 7, superiores tres laeves, convexiusculi, circa medium carinati, caeteri gradati,
convexiusculi, longitudinaliter costati, costis laevibus, crassiusculis, quam interstitiis latioribus, in
interstitiis trasversim striati, ultimus circa basim fortiter sulcatus et nodose liratus, cauda alba
oblique suleata instructus; apertura intus alba, longit. totius } haud aequans; labrum extra valde
incrassatum, album, maculis duabus fuscis ornatum, intus tenuiter liratum; columella arcuata, callo
tenui, antice denticulum formante, induta.

Longit. 8mm., diam. 4.

This is a shining semipellucid species, rather strongly costate for its size. The last three or
four ribs behind the labrum are obsolete inferiorly, and all are subnodose below the suture, through
being crossed by a shallow groove. The colour-markings are irregularly disposed, but the principal
spots are upon the upper extremities of some of the costae and at the middle and base of the
body-whorl.

104. Nassa disparilis, n. sp. (Pl. XXXYV. fig. 12). V.

Testa minima, ovata, alba, lineis rufis duabus circa anfract. ultimum cincta; anfractus 7—8,
superiores tres convexiusculi, laeves, politi, cirea medium carina filiformi instructi, caeteri longi-
tudinaliter oblique costati (costis in anfr. penult. et antepenult. crassis, circiter 8, in ultimo
gracilioribus, circiter 16, supra nodosis) inter costas spiraliter distincte suleati; apertura parva, longit.
totius 2 adaequans; labrum extra varice albo crasso transversim striato instructum, intus liris 9,
haud ad marginem productis, munitum; columella callo circumscripto induta, arcuata, transversim
tuberculata.

Longit. 7} mm., diam. 4.

Only a single example is in the collection. It is remarkable for the stoutness and small number
of the costae upon the penultimate and antepenultimate whorls in comparison with those upon the
last. The upper extremities are cut across by a distinct groove, thus forming a row of nodules

78—2

608 EDGAR A. SMITH.

below the suture. Of the two reddish lines upon the last whorl, the upper one is around the
middle, and the other at the base.

105. WNassa ecstilba, Melvill and Standen, 28,a. XIV, XV.

Family COLUMBELLIDAE.
106. Columbella conspersa, Gaskoin, 37, X/._ II.
A single typical specimen.
107. Columbella flava, Bruguiére, 37, Y7. I, X.
108. Columbella jaspidea, Sowerby, 37, X/7. 40,6,/. IX.

Only a single specimen was obtained. Probably C. plicaria, Montrouzier (Jowrn. Conch. vol. x,
p. 234, pl. rx, fig. 3, 1862) is a variety of this species. The specimen figured by Reeve is not
mottled or spotted like the type, the latter agreeing in this respect with C. plicaria.

109. Columbella galaxias, Reeve, 37, X/. XV.
110. Columbella turturina, Lamarck, 37, XZ. IX, XV.
111. Columbella varians, Sowerby, var. 40,b,/. 37, XJ. IX.

A single whitish example with the transverse striae indistinct so that the surface is rather
smooth.

Family MURICIDAE.
112. Murex ternispina, Lamarck, 37, /7//. I, VI, X, XIII.
113. Murex haustellum, Linn. 37, ///. VI, X.
114. Murex (Pteronotus) tripterus, Born. 37,///. V.
115. Murex (Chicoreus) aculeatus, Lamarck, 37,///. V, XII.
116. Murex (Chicoreus) adustus, Lamarck, 37,///. V, VII.
117. Murex (Chicoreus) rubiginosus, Reeve, 37,///. IX.
118. Murex (Chicoreus) ramosus, Linn. 37,///. IX.

119. Murex (Ocinebra) salmoneus, Melvill and Standen, 28,6. (Murex pumilus, A. Adams,
1,6. 40,6,/V. 39,9.) XII.

The name pwmilus, A. Adams, was preoccupied by Broderip many years previously, hence the
employment of the name sa/monews. I think there is little doubt that Messrs Melvill and Standen
redescribed this species, for examination of the types of both forms shows that they are the same
species. The single example from the Maldive Islands is dirty whitish, but, being a dead shell,
and occupied by a Pagurus, it probably has lost the usual rose-colour.

120. Murex (Ocinebra) pleurotomoides, Reeve, 37,///. II.

Three specimens rather smaller than the type, with the anterior canal stained with brown
and a pale brown zone round the middle of the body-whorl. The lirae within the labrum are
about six in number, and the tubercles on the columella, three or four.

The posterior sinus, at
the upper end of the outer lip, seems to be a constant feature.

MARINE MOLLUSGCA. 609

121. Murex (Ocinebra) submissus, n. sp. (Pl. XXXV. fig. 13). XII.

Testa minima, ovata, rimata, supra acuminata, albida, lineis interruptis nigro-fuscis cincta ;
anfractus 64, superiores tres laeves, convexi, caeteri supra concave declives, costis longitudinalibus
octo et liris spiralibus supra costas elatis (in anfr. penult. 2, in ultimo 8—9) instructi, incrementi
lineis tenuissimis striati, ultimus in medio convexiusculus, antice contractus; apertura intus flayescens f
labrum extra incrassatum, intus liris brevibus sex haud ad marginem productis munitum; columella
sinuosa, callo tenui reflexo flavescente induta, in medio prominens, tuberculis parvis 3—4 munita;
canalis anterior obliquus, recurvus.

Longit. 8}mm., diam. 5.

The dark brown transverse lines, about six or seven upon the last whorl, are interrupted
between the costae, and falling upon the raised lirae, the ribs appear to be crossed by coloured
tubercles.

The costae are a little oblique and continuous up the spire. The smooth protoconch is marked
off from the first normal whorl by a slender varix or thickened lip of its last whorl. The three
whorls of which it consists have a yellowish margination beneath the suture.

122. Purpura armigera, Lamarck, 37, ///. IX.
123. Purpura bitubercularis, Lamarck, 37,777. IX.
124. Purpura hippocastanum, Lamarck, 37,///. XI, XV.
125. Purpura pica, Blainville, 37, 7/7. IX.
126. LIopas situla, Reeve (Buccinum situla, Reeve, 37, 77/.) IX.
127. LIopas sertum, Bruguitre (Buccinwm sertoum, Reeve, 37,///.) I.
128, Sistrum digitatum, Lamarck (var. Jlobatum, Blainville; Ricinula digitata, Reeve,
37,/11.) IX.
129. Sistrum horridum, Lamarck (Hicinula horrida, Reeve, 37,Z//.) I, IX.
130. Sistrum ricinus, Linn. (Ricinula arachnoides, Reeve, 37, 7/1.) I, IX, XI, XV.
131. Sistrum tuberculatum, Blainville (Ricinula tuberculata, Reeve, 37,I/I.) IX.
132. Sistrum undatum, Chemnitz (Ricinula fiscellum, Reeve, 37,111. Sistrum undatum,
partim, Tryon, 41,//.) VIII, X, XV.
133. Sistrum elatum, Blainville (Ricinula spectrum, Reeve, 37,777.) II, IX, XI.
In the two fresh specimens obtained, the labrum is white, but the interior of the aperture is

yellowish.

134. Sistrum spinosum, A. Adams (Licinula chrysostoma, partim, Reeve, 37, 7/7. Ricinula
biconica, partim, Tryon, 41, //.) XIII.

Messrs H. and A. Adams (Genera Recent Mollusca, vol. 1. p. 130) recognised that Reeve had
wrongly identified the Aicinula chrysostoma of Deshayes and imposed the name of spinosws upon
the misunderstood species. The term spinosus is applicable to fig. 126 rather than to 12a, which
represents the Ricinula biconica of Blainville. I think therefore the species spinosws should be
restricted to the figure 126. Whether it is conspecific with S. biconicwm has yet to be proved.
Although united by Tryon, at present I am inclined to consider them sufficiently distinct.

610 EDGAR A. SMITH.

135. Sistrum biconicum, Blainville (Purpwra biconica, Kiener, 238. Sistrum biconicum,
Tryon, 41,72. Ricinula chrysostoma, partim, Reeve, 37,/7//.) XIII.

136. Sistrum cavernosum, Reeve (Ricinula cavernosa, Reeve, 37,/7//.) IX, XV.

137. Sistrum iostoma, A. Adams (Pl. XXXV. fig. 14). (Murex ‘ostomus, A. Adams,
cy) WHE

This species is allied to S. spinoswm, but appears to be separable.
138. Sistrum squamosum, Pease, 31,a. XI.
The Maldive specimen is quite typical.

139. Sistrum concatenatum, Lamarck (Ricinula concatenata, Reeve, 37, [II., var.= Purpura
fragum, Blainville, 23). IX.

A very variable species in size, form, and colour. Only a single small specimen was obtained
at the above station and it most resembles the variety fragwm as the aperture and labrum are
white and not orange red as in the type.

140. Sistrum decussatum, Reeve, var. (Murex decussatus, Reeve, 37,///.) XIV.

A single very small specimen differing from the type in having a shorter anterior canal, and in
the nodules at the upper angle of the whorls being uncoloured and acute. Although adult, only
13mm. in length.

Family CORALLIOPHILIDAE.

141. Coralliophila monodonta, Quoy and Gaimard (Purpura monodonta, 23). II, V, IX,
eXCe ee XC

142. Coralliophila neritoidea, Gmelin (Purpura violacea, Kiener, 37, ///.) VIII, IX.

143. Coralliophila suturalis, A. Adams, 1,6 (C. galea (partim), 41,//. Khizocheilus exu-
ratus, Pease, 31,6). IX, XV.

144. Coralliophila squamosissima, Smith, 41,//. IX.

145. Leptoconchus ellipticus, Sowerby (Magilus ellipticus, Sowerby, 40,a, XVIII.) XY.

The transverse ribbing usually extends over the greater part of the body-whorl, although, in
the description, Mr Sowerby observes that the shell is “spirally slightly ribbed below the middle,”
as if this were a specific feature. A fine example in the British Museum from the Mauritius is
27mm. long, and 22 in diameter.

146. Leptoconchus cumingii, Deshayes, 12, a, 40,a, XV/JJ. XV.

The shell figured by Sowerby (/.c. fig. 10) as Magilus globulosus, I regard as a form of the
present species.

Family LOTORIIDAE.
147. Lotorium chlorostoma, Lamarck (Zvriton chlorostoma, Reeve, 37, //.) IX, XV.
148. Lotorium gemmatum, Reeve (Zriton gemimatum, Reeve, 37, //.) IX.
149. Lotorium pyrum, Lamarck (7riton pyrum, Reeve, 37, //.) IX.
150. Lotorium tuberosum, Lamarck (Zriton tuberosus, Reeve, 37,//.) I.

151. Colubraria ceylonensis, Sowerby (Z’riton ceylonensis, Reeve, 37,//.) V, XIV.

MARINE MOLLUSCA, 611

152. Colubraria obscura, Reeve (Z'riton obscurus, Reeve, 37,//.) XIV.

153. Colubraria bracteata, Hinds (7riton bracteatus, Reeve, 37,77.) IX.

A small form only about 13mm, in length.

154, Colubraria digitale, Reeve (Z’riton digitale, Reeve, 87,1.) IX.

In fresh specimens three rows of pale brownish spots are visible upon the last whorl, one
below the suture, one at the middle, and the third below it. These spots form interrupted colour
bands, which, however, are only observable in well preserved shells.

155. Distortrix anus, Lamarck (J'riton anus, Reeve, 37,I/.) IX.
156. Ranella bufonia, Lamarck, 37,//. IX, XV.

157. Ranella granifera, Lamarck, 37,//. IX, XI, XIV.

158. Ranella tuberosissima, Reeve, 37,//. XIV.

159. Ranella thomae, d’Orbigny, 14 (A. bergeri, Canefri, 7). XII.

The single Maldive specimen has the transverse ridges of a blackish brown colour but in other
respects agrees very well with the examples cited from the above localities.

160. Ranella (Lampas) lampas, Lamarck (7’riton lampas, Reeve, 37,11.) IX.

Family CASSIDIDAE.
161. Cassis cornuta, Linn. 37,V. IX.
162. Cassis rufa, Linn. 37, V. IX.

Family DOLIIDAE.
163. Malea pomum, Linn. (Doliwm pomum, Reeve, 37, V.) IX.
164. Dolium costatum, Menke, 37,V. IX.
165. Dolium olearium, Lamarck, 37,V. XV.

166. Pirula ficoides, Lamk. jun.? 23 (Ficula reticulata, Reeve (non Lamarck), 37, J//.)
VE) VELL:

Family CYPRAEIDAE.

167, Cypraea arabica, Linn. (a. var. typica, 40,b,/V. IX. 06. var. eglantina, Duclos,
40,5,/V. IX. c. var. histrio, Meuschen, 40,),7V. IX, XI, XII, XIII. d. var. intermedia, Gray,
40,),7V. IX.).

168. Cypraea argus, Linn. 37,///. IX.

169. Cypraea asellus, Linn. 37, ///. IX.

170. Cypraea caput-serpentis, Linn. 37,///. IX, XI, XV.
171. Cypraea carneola, Linn. 37,///. I, IX, XI.

172. Cypraea caurica, Linn. 37,///. IX, XI, XIII.

612 EDGAR A. SMITH.

173. Cypraea clandestina, Linn. 37, //7. IX.

174. Cypraea cribraria, Linn. 37, ///. XIV.

175. Cypraea erosa, Linn. 37,//7. IX, XI, XIII, XV.
176. Cypraea felina, Gmelin, var. 37,777. IX, XV.
177. Cypraea gangrenosa, Solander, 40,0,/V. X, XII.
178. Cypraea (Epona) globulus, Linn. 37,///, XIII.
179. Cypraea helvola, Linn. 37, //7. VIII, IX.

180. Cypraea hirundo, Linn. 37,///. IX, XIII.

181. Cypraea isabella, Linn. 37,//7/. IX, XI.

182. Cypraea lamarckii, Gray, 37,///. III, VIII.
183. Cypraea lynx, Linn. 37,///. I, IX.

184. Cypraea mauritiana, Linn. 37,///. XI.

185. Cypraea minoridens, Melvill, 27,1, 39,d (Cypraea fimbriata, partim, Sowerby,
40,6,/V.) IX.

186. Cypraea moneta, Linn. 37,///7. IX, XI, XIII, XIV, XV.

187. Cypraea poraria, Linn. 37,///. IX, XI.

188. Cypraea annulus, Linn. 37,///. IX.

189. Cypraea staphylaea, Linn. 37,///. VII.

190. Cypraea talpa, Linn. 37,///. IX.

191. Cypraea tigris, Linn. 37,///. IX.

192. Cypraea vitellus, Linn. 37,///. IX.

193. Cypraea walkeri, Gray, 37,///. II, III, V, VI, XIII.

194. Cypraea ziczac, Linn. 37,///. XIII.

195. ‘Trivia pellucidula, Gaskoin (Cypraea pellucidula, Sowerby, 40,0,/V.) X.

Family STROMBIDAE.
196. Strombus dentatus, Linn. (var. Riippellii, Reeve, 37, VI.) I—VI.
197. Strombus floridus, Lamk. 37, VJ. I, IX, XI, XV.
198. Strombus gibberulus, Linn. 37, V/. IX—XII, XIV, XV.
199. Strombus labiosus, Gray, 37, VJ. V, VI.
200. Strombus lentiginosus, Linn. 37, V/. IX.
201. Strombus guttatus (Chemnitz), Kiener, 23, 37, VJ. V.
202. Strombus lamarckii, Gray, 40,),/. IX.

MARINE MOLLUSCA. 613

203. Pterocera lambis, Linn. 37, V/. IX.
204. Pterocera chiragra, Linn. 37, V/. IX.
205. Terebellum terebellum, Linn. (Terebellum punctatum, Reeve, 37, VI.) II—VI, X.

Family CERITHITDAE.

206. Triforis corrugatus, Hinds, 20,a,8. X.

207. Triforis elegans, Hinds, 20, a (7. pictwratus, Sowerby, 40,¢). IX. '

I have compared the type of 7. pictwratus with authentic specimens of the present species
and have no hesitation in pronouncing them specifically the same. Although Mr Sowerby described
his shell as having three keels on a whorl, there are in fact four, and they alternate, larger and
smaller, as stated by Hinds. No mention is made by either author of the spotted carina below
the periphery of the body-whorl. Triforis is a genus which wants monographing by a competent
person, and ought to be illustrated with much enlarged photographed figures. At present it is
practically impossible to identify most of the described species by means of the published figures
and descriptions.

208. Triforis sculptus, Hinds, 20,a. XIV.

This species varies much in size, and is sometimes considerably larger than the shell figured
by Hinds or the dimensions given in his original description’. The largest specimen in the British
Museum is 20mm. in length, whereas another, equally adult, is only 15.

209. Triforis lilaceocinctus, n. sp. (Pl. XXXYV. fig. 15). XIV.

Testa elongato-pupoidea, supra acuminata, granulis flavescentibus et albis, lirata, inter granula
rufo punctata; anfractus circiter 15, superiores liris duabus granosis aequalibus instructi, pauci
anteriores lira graciliore mediana, dilute lilacea, vix granulata ornata, sutura lineari sejuncti, micro-
scopice spiraliter striati, ultimus circa basim lilaceus, liris aliis tribus nodulosis, rufo punctatis,
instructus; apertura obliqua, piriformis ; canalis parvus, dextrorsus, semiclausus.

Longit. 10} mm., diam. 3} mm.

A general glance at this pretty species gives the impression that the whorls have each two
adjacent rows of large granules, Such however is not the case. It is the lower row in one whorl
being adjacent to the upper one in another (the linear suture being between) that gives this
appearance, the unspotted and more slender median lirae also lending to the effect. This lira
gradually dies out as it ascends the spire, so that the upper whorls have only two rows of equal
sized granules. This species also occurs at the Mauritius (Brit. Mus.).

210. Triforis excellens, n. sp. (Pl. XXXYV. figs. 16, 17). V.

Testa magna, dilute fuscescens, hic illic saturatius maculata; anfractus numerosi (25—30), plani,
cingulis tribus aequalibus acutis cincti, inter cingula laeves, ultimus ad peripheriam bicingulatus,
infra vix convexus, liris 4—5 instructus; apertura irregulariter quadrata; labrum tenue; columella
callo reflexo crassiusculo induta.

Longit. 29 mm., diam. 6 mm.

The spirals are equal in size, but the space separating the lowermost from the median one
is a little broader than the groove between the upper two. The suture except at the upper part
of the spire is marked with a spiral thread,

1 Ann. Mag. Nat. Hist. 1843, vol. x1. p, 17.

614 EDGAR A. SMITH.

211. Triforis gracilior, n. sp. (Pl. XX XV. figs, 18, 19). V.

Testa 7’. excellenti similis, sed gracilior, cingulis duobus inferioribus subundulatis, alba, hic illic
fusco maculata,

Longit. 28mm., diam. 54mm.

Like 7. excellens, this species has three keels upon each whorl, but they are not so equal,
the uppermost being a trifle more slender than the others. The latter also are slightly affected
by faint longitudinal depressions giving them an obscurely beaded appearance. The body-whorl is
bicarinate at the periphery and has three or four lirae beneath. As in the preceding species, the
space between the second and third keel is a little broader than that which separates the first
and second, and the suture is filo-lirate.

212. Triforis pura, n. sp. (Pl. XX XV. figs. 20, 21). XII.

Testa elongata, alba, clathrata, granulata; anfractus circiter 20, fere plani, costis spiralibus
tribus granosis inaequalibus (costa mediana minima) cincti, inter costas longitudinaliter costati,
ultimus costis senis (prope labrum duabus intercalatis) ornatus; canalis dextrorsus, clausus, laevis ;
apertura irregulariter ovata, obliqua; labrum antice prominens, postice recedens; columella callo
erassiusculo reflexo induta.

Longit. 14mm., diam. 3.

Of the three spirals the uppermost is a little stouter than the lowermost and the central one
is rather finer than the latter. The granules form oblique rows of three, being connected by the
longitudinal costae. The suture is thread-like.

213. Wertagus fasciatus, Bruguiére, var. 40,a,XV. VIII, IX.
214. Wertagus kochi, Philippi, 40,0, YV. XI.

215. Wertagus articulatus, A. Adams and Reeve (Cerithium articulatum, Adams and Reeve,
3. V. articulatus, Sowerby, 40,a,XV.) VI, X.

216. Wertagus obeliscus, Bruguiére, var. cedo-nulli (V. cedo-nulli, Sowerby, 40, a, XV.)
IL, IDS,

V. cedo-nulli is perhaps separable from obeliscus.

217. Cerithium columna, Sowerby, 40,0, XV. IX, XI, XIII.

218. Cerithium echinatum, Lamarck, 40,a, XV. XIII.

219. Cerithium lacteum, Kiener, 40,a,XV. XI.

220. Cerithium piperitum, Sowerby, 40,a,XV. IX.

221. Cerithium salebrosum, Sowerby, 40,a,XV. II.

222. Cerithium tuberculatum, Linn. (var.=(. variegatuwm, Quoy and Gaimard, 40, a, XV.) XI.

223. Cerithium albovaricosum, Smith, var. 39,9. I.

Three specimens, a little larger (19mm. long), paler, and with less of the brown dotting between
the costae than in the type.

224. Terebralia palustris, Linn. (Pyrazus palustris, Sowerby, 40,a,XV.) XI.

225. Modulus tectum, Gmelin, 41,/X. IX.

MARINE MOLLUSCA. 615

Family PLANAXIDAE.

226. Planaxis ineptus, Gould, 17, 40,a, XX, 41,/X. X.

The Maldive specimens are a little larger than the dimensions given by Gould, being 5} mm.
in length. Most of them also exhibit a purplish stain on the columella which is not referred to
in the original description. There are generally three principal colour lines upon the body-whorl
—one above the periphery, one below it and the third still more anterior.

Family LITTORINIDAE.
227. Littorina glabrata, Philippi, 33,0, 37,X. XI, XV.
228, Littorina scabra, Linn. 37, X. XV.
229. Littorina undulata, Gray, 37, XY. XI, XV.

Family SOLARITDAE.
230. Solarium modestum, Philippi, 40,0,/. V.
231. Philippia cingulum, Kiener, 40,0}, ///. II, IX.

Family HIPPONYCIDAE.
232. Mitrularia cicatricosa, Reeve (Calyptraea cicutricosa, Reeve, 37, XJ.) IX.
233. Mitrularia equestris, Linn, (Calyptraca equestris, Reeve, 37, XJ.) IX.
234. Hipponyx danieli, Crosse (Capulus danieli, Crosse, 11,6). ILI—V.
235. Hipponyx australis, Lamarck, 41, V///.

; Family CAPULIDAE.
236. Thyca crystallina, Gould (Hipponyx crystallinus, Tryon, 41, ViZ/.) VIII.

Family XENOPHORIDAE.
237. Xenophora indica, Gmelin, 41, ////. (Phorus indica, Reeve, 37,7.) V.

Family VANICOROIDAE.
238. Wanicoro cancellata, Lamarck, 41,V///., 40,6,V. IX, XIII.

Family NATICIDAE.
239. INatica robillardi, Sowerby, 40,d. XII, XV.

240. Natica euzona, Recluz (Pl. XXXV. fig. 22), 36 (WV. picta, partim, Reeve, 37, 7X.
NV. decora, Philippi, 33,a). VII.
The operculum of this species is shelly, white, concave externally, shining except in the middle

where there is a dull opaque thickening. The outer curved side has a thickened edge, within

79—2

616 EDGAR A. SMITH.

which is a parallel furrow and then a raised ridge or rounded keel. The convex side is covered
with a thin yellow pellicle and is marked with lines of growth.

241. Polinices mamilla, Lamarck (Natica mamilla, Reeve, 37,/X.) III, IX, XII, XV.
242. Polinices albumen, Lamarck (Watica albumen, Reeve, 37,7X.) IL
243. Mamilla melanostoma, Lamarck (Watica melanostoma, Reeve, 37,7X.) VI, XV.

244, Mamilla simiae, Deshayes (Natica simiae, Reeve, 37,/X.) IX.

Family IANTHINIDAE.

245. Ianthina communis, Lamarck? 37, X/, IX.

The figures most resembling the specimens from the above station are Reeve’s figures of
I. affinis (figs. 2 a,b). That so-called species and several others described in the same monograph
are probably mere varieties of one variable species. (See Tryon, Man. Conch. vol. 1x. pp. 36, 37.)

Family SCALIDAE.

246. Scala replicata, Sowerby (Scalaria replicata, Sowerby, 40, 6,7.) IX.

247. Scala consors, Crosse and Fischer (Scalaria consors, Tryon, 41, 7X.) XI.

A single example considerably larger than the type, being 18mm. long, and 7 mm. in width.
The locality given by Crosse and Fischer is St Vincent’s Gulf, 8S. Australia, but four specimens,
supposed to include the type among them, were presented to the British Museum in 1870 by
Mr G. F. Angas and were marked “Ceylon” by him. The Maldive example has a dark line at
the suture.

248. Scala (Cirsotrema) texta, n. sp. (Pl. XXXV. fig. 23). X.

Testa elongata, subulata, imperforata, solidiuscula, albida, longitudinaliter plus minus plicata,
varicibus paucis obliquis instructa, spiraliter suleata et tenuiter punctato-striata, lineis incrementi
tenuissimis obliquis sculpta; anfractus circiter 12, supra valde declives, vix concavi, in medio obtuse
angulati, sutura obliqua irregulariter corrugata sejuncti, ultimus ad peripheriam angulatus, costis ad
angulos subnodosis; apertura oblique ovata; peristoma continuum, expansum, margine externo varice
crasso circumdato. Longit. 17 mm., diam. 5; apertura intus 3 longa, 2 lata.

This very distinct species is well-characterized by the fine cancellated sculpture, like a piece
of linen, the interstices being minutely punctate. The whorls exhibit spiral shallow sulci and
intervening raised ridges, both the latter and the grooves being covered with the spiral and longi-
tudinal striae. Every second or third whorl exhibits an oblique varix, and the upper ends of the
plicae form a very irregular puckered or dentate sutural line.

Allied to S. bicarinata, Sowerby, but much larger, of different form, differently plicate and
spirally sulcate.

Family PYRAMIDELLIDAE.
249, Pyramidella corrugata, Lamarck, 23. XV.
250, Pyramidella nodicincta, A. Adams, 40,b,//. XIV.

251, Obeliscus dolabratus, Linn. (0. dolabratus and O. terebellwm,° 40,6, II.) IX, XV.

MARINE MOLLUSCA. 617

252. Obeliscus monilis, A. Adams, 40,0, //. IX, XIV.
253. Obeliscus sulcatus, A. Adams, 40,),//. XV.

Family NERITIDAE.
254. WNerita albicilla, Linn. 37,/X. IX, XV.
255. WNerita histrio, Linn. 37,/X. XI.
256. WNerita plicata, Linn, 37,/X. IX, XI.
257. Werita polita, Linn. 37,/X. I, XII.

Family TURBINIDAE,

258. Turbo (Senectus) argyrostoma, Linn. 37, /V. (var.= 7. margaritaceus, Reeve, 37, IV.)
EXC OXGT EXGV),

Family LIOTIIDAE.

259. lLiotia squamicostata, n. sp. (Pl. XXXV. fig. 24). X.

Testa subglobosa, anguste umbilicata, alba; anfractus quatuor, ad suturam canaliculati, costis
granosis spiralibus (in anfractu penultimo 4, in ultimo 8) cincti; granula ad peripheriam subsqua-
miformia; apertura circularis, margine externo crenulo, columellari incrassato, reflexo, Diam. maj.
5 mm,, alt. 4.

The spiral ridges are rather strong, broader than the grooves between them, and the granules
upon the second, third, fourth, and fifth upon the body-whorl, and the second and third upon the
penultimate, are scale-like, and that which encircles the umbilicus is much thicker than the rest and
transversely grooved. The terminations of the spirals upon the outer lip give it a crenulated aspect.

Family TROCHIDAE.

260. Trochus (Lamprostoma) incrassatus, Lamarck, 34, X¥/. I, XV.

261. Trochus (Lamprostoma) maculatus, Linn. 34, YJ. IX, XI.

262. Trochus (Lamprostoma) obesus, Reeve, 37, X///J. IX.

263. ‘Trochus (Lamprostoma) radiatus, Gmelin, 34, X/. XI.

264. Thalotia maldivensis, n. sp. (Pl. XXXV. figs. 25, 26). II, X.

Testa parva, elate conica, imperforata, ad peripheriam acute angulata, supra viridis, ad suturam
albo et roseo maculata, apicem versus rubescens, supra anfractum ultimum strigis albis obliquis paucis
ornata, infra roseo et albo tessellata; anfractus 8—9 plani, seriebus granulorum parvorum septenis
(quarum series duae inferiores contiguae et albo et roseo maculatae) instructi, ultimus infra angulum
seriebus 9—10 ornatus, subplanus; apertura obliqua, subquadrata, margaritacea, iridescens; labrum
tenue; columella alba, incrassata, reflexa, leviter arcuata, antice truncata.

7

Diam. maj. 7} mm., min. 7, alt. 10. Var. Testa rufescens, concolor.

A very pretty conical shell, generally more or less green with a white and pink spotted band

above the suture and dotted with pink in the middle of the base. Some examples are uniformly
pinkish reddish and with. only tinges of green above or below.

618 EDGAR A. SMITH.

265. Clanculus atropurpureus, Gould (Zrochus (Clanculus) atropurpureus, 34, XI.) 1X, XIII.

266. Perrinia angulifera, A. Adams (Pl. XXXV. fig. 27); (urcica (Perrinia) angulifera,
34, XJ). X.

267. Forskalia pulcherrima, A. Adams (Gibbula pulcherrima, Pilsbry, 34, XZ.) IV, VI.
268. Monilea calyculus, Wood, 34,X/. II—IV.

269. Monilia simulans, Smith, var. 39,c. V, VI, X.

These specimens differ from the type in having only very faint indications of the nodose
plications at the upper angle of the whorls. They also differ in colour, being paler and less rosy.
Trochus (Monilea) warnefordi, G. and H. Nevill, may be the same as this species.

270. Ethalia rhodomphala, n. sp. (Pl. XXXVI. figs. 1, 2). III, V, X.

Testa orbiculata, supra breviter conica, semiobtecte anguste umbilicata, ad peripheriam acute
angulata, supra albida, maculis paucis remotis fuscis et lineis tenuissimis irregularibus undulatis
radiatim picta, ad peripheriam maculis semipellucidis roseo pictis ornata, infra lineis albis opacis
irregularibus subzigzagformibus variegata, circa umbilicum rosea; anfractus sex leves, nitidi, ultimus
circa medium striis paucis sculptus, circa umbilicum radiatim plicatus; apertura oblique subquadrata,
margaritacea, iridescens; columella crassa, in medio margaritacea, valde reflexa, alba vel rosea, callosa,
umbilicum semiobtegens.

Diam. maj. 8 mm., min. 7, alt. 33.

?

Viewed with the naked eye, the upper surface of this shell looks whitish with a few brown
radiating blotches. A closer inspection with a lens, however, reveals the presence of very fine
lineation, more or less broken up, undulating or dotted. The puckering around the umbilicus is
pinkish, the rest of the base being covered with opaque white irregular markings upon a sub-
translucent ground. The last whorl is angled at the periphery and above the angle is marked
with a few distinct spiral striae.

Family STOMATELLIDAE.
271. Gena auricula, Lamarck, 34, Y/7. IX, XIII.

Family HALIOTIDAE.
272. Haliotis ovina, Chemnitz, 34, X77. IX, XII.

The number of open perforations may vary from two to six.

Family FISSURELLIDAE.
3. Glyphis bombayana, Sowerby, 34, Y//. IX.
274. Glyphis singaporensis, Reeve, 34, Y//. XIII.
5. Emarginula incisura, A. Adams, 34, Y//. XIII.

The locality of this species has not previously been recorded. The single specimen obtained
is of a greenish yellow tint externally and the central portion of the interior is of a somewhat
similar colour. The anterior slit is very peculiar, being clean-cut, and extending from the margin
almost to the apex.

MARINE MOLLUSCA. 619

276. Emarginula planulata, A. Adams, 34, X//. VIII.

A single specimen apparently inseparable from this species.

Family CHITONIDAE,
277. Chiton (Sclerochiton) miles, Carpenter, 34, X/V. I.

Torres Straits is the only locality previously known for this Chiton.

278. Schizochiton incisus, Sowerby, 34, X/V. I.

This I believe is the first record of the occurrence of this remarkable species in the Indian Ocean,

Family ISCHNOCHITONIDAE.
279. Callochiton platessa, Gould, 34,X/V. V.
280. Ischnochiton hululensis, n. sp. (Pl. XXXVI. figs. 3—6). IX.

Testa ovata, mediocriter elata, roseo-albida, punctis roseo-griseis numerosis ornata, cingulo minute
squamulato, albido viridi-griseo maculato, circumdata; valvae undique minute decussatae, antica
transversim corrugata, lamina inserta brevi, fissuris novem instructa, medianae areis lateralibus bene
circumscriptis, elatis, transversim corrugatis, postica mucrone centrali instructa, dimidio posteriori
convexo, anteriore concavo et corrugato, lamina inserta fissuris undecim parata; squamae cinguli
minutae, acute ovales, transversim striatae.

Longit. 13 mm., diam. 19.

The single specimen obtained is of a pale pinkish colour down the sides of the valves and
whitish flecked and spotted with greenish grey down the middle. The hinder almost straight edge
of the valves is pale with a few greenish grey spots and a dash of the same colour occurs along
the sides in front of the lateral areas. The girdle is covered with alternate patches of greyish
green and white scales. The insertion-plates of median valves have a single slit, that of the front
valve has nine slits and that of the tail-valve eleven.

281. Ischnochiton feliduensis, n. sp. (Pl. XXXVI. figs. 11—14). VII.

Testa ovata, mediocriter elata, in medio carinata, pallida, sordide alba; valvae tenuissimae,
antica et areae laterales valvarum centralium minute granulatae; areae centrales longitudinaliter
grano-liratae ; valva anterior semicircularis, postice in medio sinuata; mucro valvae posterioris ante-
medianus, parum prominens; cingulum album, squamis minutis striatis ornatum. Longit. 6 mm.,
diam. 4mm.

Remarkable for the thinness of the valves, the flatness of the lateral areas, and its style of
sculpture. The posterior valve has twelve slits in the insertion plate, the anterior has nine and
the intermediate valves one.

282. Ischnochiton maldivensis, n. sp. (Pl. XXXVI. figs. 7—10). VII.
Testa elongato-ovata, elata, convexa, in medio subcarinata, pallida, plus minus rufescens, albo
minute punctata, cingulo minute squamulato griseo maculato circumdata; valva antica et areae

laterales laeves, areae centrales sulcis paucis longitudinalibus utrinque sculptis, valya posterior in
medio mucronata; squamae cinguli minutae, adamantiformes, indistincte striatae.

Longit. 8mm., diam. 4.

The salient features of this species are the smoothness of the head-valve and the lateral areas,
and the distinct imcised lines upon the sides of the central areas, a space down the middle. of

620 EDGAR A. SMITH.

the shell being smooth. The insertion plates are thick and striated across the edge. The anterior
valve exhibits eight slits, the posterior ten and the median valves none.

Family ACANTHOCHITIDAE.

283. Acanthochites (Loboplax) laqueatus, Sowerby, 34, X/V. (Angasia tetrica, 34, XI 4)
WoC, DING

Testa parva, elongata, subdepressa, purpureo-rosea, albo variegata, valvis antica et postica
pallidis, cingulo puniceo vel viridi, hic illic albo maculato, cireumdata; valvae rugose squamatae ;
antica costis radiantibus quinque vix elatis instructa, undique squamata, medianae angustae, in
medio postice rostratae, area dorsali sublaevi, postica undique squamata, breviter mucronata,

Longit. 16 mm., diam, 12mm.

This species is remarkable for its brilliant variable colour, the narrowness of the central valves
and their coarse squamulation.

The anterior valve in the Maldive specimens is greenish grey with five paler rays and the
posterior margins are also whitish with one or two black spots. Central valves purplish rose with
some whitish streaks or wrinkles on each side the dorsal area, Mantle purple red with a white
spot on each side the first, seventh and eighth valves surrounding the lateral pores there situated,
also some black spots set in white rings all round the edge, about 17 or 18 altogether, and with
a fringe at the extreme margin of short white spicules.

Another specimen has the valves similarly coloured, only the seventh is decidedly green
excepting the rosy-purplish dorsal area. The mantle, however, is of a vivid green, with a white
stripe on each side the anterior valve and four or five stripes or spots in front, the outer margin
with a similar fringe of spicules.

Valves. First valve semicircular, the posterior margins being slightly divergent, exhibiting five
not much raised rays which, like the rest of the surface, are covered with coarse elongate scales
which are finer towards the middle of the hinder edge. Insertion plate finely striated externally,
acute at the edge, with five shallow slits.

Central valves narrow, curved in front and rather acutely beaked behind; dorsal area narrow,
smooth, with cross marks of growth; squamae arranged in rows, some parallel with the posterior
margin, others obliquely across the valves, three or four rows on each side the dorsal area being
white; insertion plates with a single posterior slit.

Posterior valve semicircular, shortly mucronate, squamate everywhere, except upon the short
mucro which of course is like the dorsal area of the preceding valve; insertion plate thickish,
with nine slits.

The colour of the interior of the valves of the only specimens disarticulated is a pinkish flesh
tint, the head and tail valves being*paler than the rest.

Tt does not seem to me advisable to place this species in the genus Angasia as distinct from
Acanthochites. I might also point out that the name Angasia was preoccupied in Crustacea. (See
P. Z, 8S. 1863.)

Family CRYPTOPLACIDAE.

284, Cryptoplax burrowi, Smith, 34,XV. IX.

MARINE MOLLUSCA. 621

Family NEOMENIIDAE.

285. Proneomenia australis, Thiele, 34, XVJ/. V.

Only a single specimen, apparently belonging to this species, was obtained.

Family SPONDYLIDAE.
286. Spondylus imperialis, Chenu, 37,/Y. IV.

Family LIMIDAE.
287. Lima (Mantellum) fragilis, Chemnitz, 40,b,/7. I, IX, XI, XV.
288. Lima (Mantellum) inflata, Chemnitz, 40,0, 7. (Lima fasciata, 40,b,7.) XIII.
289. Lima squamosa, Lamarck, 40,6,7. II.

290. Lima (Ptenoides) tenera, Chemnitz jun.? 40,4,/. V.

: Family PECTINIDAE.
291. Pecten senatorius, Gmelin, 40,),/7., 37, V///. IJI—VI.

Only young specimens obtained. P. /ayardi, Reeve, P. cristularis, Adams and Reeve, P. rugosus,
Sowerby and some other so-called species, I regard as half-grown or young specimens of the
present form.

292. Pecten lemniscatus, Reeve, var. 37, V///._ II.

293. Pecten mirificus, Reeve, 37, V///. V.

294. Pecten noduliferus, Sowerby, 40,),7. V, XII.

295. Pecten irregularis, Sowerby, 400,/7., 15. V, X, XII.

Remarkable for the very numerous fine costae which are closely prickly scaled in fresh young
specimens.

296. Pecten nux, Reeve, 37,V///. II, V, VI.
297. Pecten cooperi, n. sp. (Pl. XXXVI. figs. 15—18). VII.

Testa parva, compressa, albida, rufo maculata, aequivalvis; valva sinistra costis principalibus
novem et aliis gracilioribus intercalatis instructa, supra costas pulcherrime concentrice lamellata ;
auriculae valde inaequales, postica minima, antica costis 2—35 magnis aliisque paucis tenuibus ornata,
transversim tenuiter lamellata, lamellis supra costas incrassatis, conspicuis; valva dextra costis
circiter 12, tri- vel quadripartitis, instructa, concentrice tenuiter lamellata; auricula antica costis
tenuibus radiantibus circiter 10 squamulatis ornata, costa suprema marginali maxima.

Longit. 124 mm., alt. 144, diam. 4.

A single example only was obtained. The species has very remarkable sculpture and very
unequal auricles. The concentric lamellae are similar on both valves but the costae are different.
On the left valve there are about nine principal ones with a secondary one between them and
often with still finer ones on each side of the secondaries. The right valve has about twelve costae
each made up of three to five small ones, or in other words it is ornamented with numerous

G. ‘II; 80

622 EDGAR A. SMITH.

slender riblets arranged in groups of three to five. Named after Mr Forster Cooper who was
associated with Mr Stanley Gardiner in the exploration of these islands.

298, Pecten maldivensis, n. sp. (Pl. XXXVI. figs. 19, 20). II, IV, VI.

Testa compressissima, albida, plus minus roseo radiata et maculata, et maculis albis opacis
interdum variegata, tenuissime radiatim striata, lineisque incrementi tenuibus sculpta; valvae sub-
aequales, dextra vix convexiore; auriculae inaequales, posticis paulo majoribus, levibus, anticis costis
tenuissimis paucis radiantibus ornatis.

Longit. 17 mm., alt. 19, diam. 33.

Very compressed and usually very finely radiately striated, but sometimes smoother, with less
distinct striae. The colour is variable, some specimens are entirely white with only a few colour-
spots on the dorsal slopes. Others are dirty whitish, with about seven rather obscure reddish rays
and one example has the right valve rayed with red and the left blotched with opaque white and
red. All exhibit the red-spotting at the sides of the dorsal slopes.

299. Janira gardineri, n. sp. (Pl. XXXVI. figs. 21, 22). V, VIL.

Testa mediocriter compressa, suborbicularis, costis 19 planatis, quadratis, latioribus quam sulci,
instructa, undique confertim tenuissime concentrice lamellatay valva sinistra plana, albida, rufo
lineata et punctata, dextra alba, umbones versus rufo tincta; umbones laeves, acuti, contigui, haud
radiatim striati, nitidi, albidi, lateribus convergentibus rectis; auriculae inaequales, posticis paulo
majoribus, costis paucis radiantibus tenuibus parum conspicuis instructae, transversim tenuissime
lamellatae, et linea cardinis valvae dextra squamicristata; pagina interna alba.

Longit. 25 mm., alt. 23, diam. 6.

A very beautiful species well characterized by the flat-topped and square-cut costae which are
rather broader than the intervening grooves. The concentric fine lamellae seem liable to disappear
from the grooves in the flat valve, but they are very conspicuous on the costae and towards the
umbo are thickened, so that they are more like granules than lamellae. The red markings occur
both on and between the ribs. In the convex valve the lamellae are as conspicuous in the sulci
as upon the costae. The lamellae upon the auricles are much more delicate and closer than those
upon the rest of the shell. This species probably attains larger dimensions than those given above.

300. Pallium plica, Linn. (Pecten plica: Reeve, 37,VZIZ). II—VI.
301. Amussium paucilirata, n. sp. (Pl. XXXVI. figs. 23, 24). V.

Testa parva, compressa, tenuis, pellucida, laevis, polita, aequivalvis, valva sinistra alba, dextra
pellucida albo et rufo radiata et variegata, radio conspicuo albo supra latus anticum ornata; auriculae
inaequales, postica valvae sinistrae infra leviter sinuata, posteriori valvae dextrae majori; umbones
acuti; pagina interna radiatim substriata, lira brevi unica utrinque instructa.

Longit. 74 mm., alt. 74, diam. 2.

An interesting little species, having one valve pure white and the other mottled and rayed
with white and red. The presence of only two internal lirae is also remarkable. In addition to
these a short lira runs from the ligament-pit along the base of each auricle. Although the
surface of both valves looks smooth and glossy, under a lens fine striae of growth are visible,
The umbones are acute, that in the coloured or right valve being a little more prominent than
that of the left. The opaque white ray down the anterior side appears to be a specific feature,
for it is present in all the specimens examined.

302. Semipecten forbesianus, Adams and Reeve, 37, V/.

MARINE MOLLUSCA.

Family PTERIIDAE.
Malleus anatinus, Gmelin, 37, X/7. IV.
Vulsella rugosa, Lamarck, 13. XII.
Electroma ala-corvi, Chemnitz (Avicula ala-corvi, Reeve, 37,X.) XV.

Electroma zebra, Reeve (Avicula zebra, Reeve, 37, X.) X.

623

Melina perna, Linn. (Ostrea perna, Hanley, 19,6. Perna (Isognomon) samoensis, Baird,

Pinna muricata (Linn.), Reeve, 37, XZ. XV.
Pinna nigra (Chemnitz), Dillwyn, 37,X/7. IX.
Pinna saccata, Linn. 37, X/7. I.

Pinna serrata, Solander (jun.?), 37, XZ. VII.

Family MYTILIDAE.
Modiola arborescens, Chemnitz, 37, XY. IX, X, XIII, XIV.
Modiola auriculata, Krauss, 25, 10. IX.
Modiola flavida, Dunker, 37, ¥. X.
Lithodomus cinnamominus, Chemnitz, 37, Y. II.
Lithodomus gracilis, Philippi, 37, X. XII.
Lithodomus malaccanus, Reeve (Young?), 37,X. IX.
Lithodomus nasutus, Philippi var.? 37,X. XIV.
Lithodomus teres, Philippi? 37, XY. VII, XV.
Modiolaria argentea, Reeve (Lithodomus argentews, Reeve, 37, X.) II.

Septifer bilocularis, Linn. (Mytilus nicobaricus, Reeve, 37,X.) XII.

Family ARCIDAE.
Arca maculata, Sowerby, 37,/7/. I, IX, XI, XV.
Arca imbricata, Bruguiére, 37,77. IX, XV.
Arca (Anadara) clathrata, Reeve, 37,//. VII.
Arca (Anadara) urypygmelana, Bory, 37,//. VII, IX.
Arca (Anadara) antiquata, Linn. 37,//. Maldives.
Arca (Scapharca) rhombea, Born, 37,//. IX.
Arca (Acar) symmetrica, Reeve, 37,77. VII, VIII.
Arca (Acar) domingensis, Lamarck, 24. II, VII, IX, XIII, XIV.

624 EDGAR A. SMITH.

330. Arca (Barbatia) lacerata, Linn. 37,//. II.
331. Arca (Barbatia) lima, Reeve, 37,//. II, V, IX, XIV, XV.
332. Arca (Barbatia) tenella, Reeve, 37,//. I, IX, XV.

Family CARDITIDAE.

333. Mytilicardia variegata, Bruguiére (Cardita variegata, Reeve, 37, 1.) I, EXE EXcIe OVE

Family GALEOMMIDAE.

334. Galeomma formosa, Deshayes, 40,),//7. XIV.
335. ‘Thyreopsis coralliophila, H. Adams, 2. II.

In this remarkable genus the mantle is united ventrally, with a small anterior slit for the
passage of a small pointed foot, and a very minute anal opening behind, near to which the
surface is somewhat scabrous or minutely papillose. There is a narrow free fold of the mantle
bordering the margin of the valves which are thickened within and gives rise to a delicate riblet
which extends across the inner surface of each valve from a point near the middle about half
way towards the umbo'. Only faint traces of adductor scars are visible and probably the free
edge of the mantle above referred to is the principal means of attachment to the shell. It does
not appear as if it would be reflected upon the exterior of the shell when living. In a form
where the valves are permanently more or less spread out almost horizontally, the animal being
too large for them to close, adductor muscles would be of little service, and therefore we might
expect to find them but feebly developed. The shell is of a purplish tint within and without,
except towards one end, which is white. The exterior is excessively finely concentrically striate
and microscopically radiately striate and the inner surface of the valves appears to be minutely
pitted. The dorsal margins are thickened within and the ligament is attached in a depression of
this thickening immediately beneath the minute, glossy, convex, boss-like umbones.

Family TRIDACNIDAE.

336. Tridacna cumingii, Reeve, 37, X/V. IX.

Family CARDIIDAE.
337. Cardium leucostoma, Born, 37,//7. IX.
338. Cardium nebulosum, Reeve, 37, //. II.
339. Cardium maculosum, Wood, 37,//. III.
340. Cardium levisulcatum, n. sp. (Pl. XXXVI. figs. 25, 26). IX.

Testa irregulariter quadrata, inaequilateralis, alba, mediocriter convexa, latere antico brevi,
rotundato, postico oblique curvatim truncato, multo majori, margine ventrali late arcuato; valvae
radiatim anguste sulcatae quam costis multo angustiores; costae planiusculae, circiter 25, quarum

1 This riblet is only feebly indicated in Adams’s type in the British Museum by a slight sinuation of the
margin at that point.

MARINE MOLLUSCA, 625

anteriores squamis transversis subgranuliformibus subconfertis instructae, posteriores haud granulatae
sed transversim tenuissime striatae.

Longit. 15 mm., alt. 12, diam. 84.

Remarkable for its somewhat compressed and quadrate form, the slender radiating sulei and
the slightly raised ribs, of which only the anterior half bear transverse granules, those on the
hinder half of the shell being only very finely transversely striated. The colour is entirely white
excepting the dorsal margin just in front of the umbones, which is stained with pink.

341. Cardium sueziense, Issel, 21, 39,0. II, X, XV.

342. Cardium (Laevicardium) lobulatum, Deshayes, 12,c. IJ, III, V, VII.
343. Cardium (Laevicardium) australe, Sowerby, 37,77. IX, XII, XIII, XV.
344. Cardium (Laevicardium) biradiatum, Bruguiére, 37, //. V, VII, IX, XII.

345. Cardium (Ctenocardia) victor, Angas, 4. VIII.

The ribs on the greater part of this beautiful shell are alternately larger and smaller, the
former being spinosely squamate and the latter only very delicately scaled, On the posterior end
the costae are about equal in stoutness and ornamented with short scale-like spines.

346, Cardium (Ctenocardia) fornicatum, Sowerby, 37,//. V, X.

Family CHAMIDAE.
347. Chama jukesi, Reeve, 37,/V. I.
348. Chama imbricata, Broderip, 37,/V. IX.

Family CYPRINIDAE.
349. Meiocardia lamarckii, Reeve, 37,//. V.
350. Trapezium oblongum, Linn. 37,/. V.

Family VENERIDAE,
351. Venus reticulata, Linn. 37,X/V. I, IX.
352. Wenus marica, Linn. 37,X/V. IX, XV.
353. Wenus recognita, Smith, 39,a. II. «
354. Wenus toreuma, Gould, 37, X///. IX.
355. Circe scripta, Linn. 37,X/V. VI.
356. Ibioconcha picta, Lamarck (Circe picta, Reeve, 37, X/V, 38.) III—V, VII, IX, X,

357. Lioconcha sulcatina, Lamarck, 38. VIII, var. IX.

The variety differs from the type in being smooth anteriorly, without the fine yet conspicuous
concentric striae which usually occur at that end of the shell. In form and colour-pattern they
exhibit scarcely any difference. One example however is entirely white.

626 EDGAR A. SMITH.

Family PETRICOLIDAE.
358. Petricola lapicida, Chemnitz, 40,a, X/X. II, VIII, XV.

Family PSAMMOBIIDAE.
359. Psammobia rugulosa, Adams and Reeve, 37,X. X.

360. Psammobia pulcherrima, Deshayes, 37, X, 39,a. X.

The Maldive specimen is considerably larger than the type described by Deshayes, being
31 millim. long, and 16 broad. It is whitish with several irregular somewhat broken purple red
rays which are also visible upon the inner surface of the valves. The scales upon the posterior
costae become obsolete as the shell increases.

361. Asaphis deflorata, Linn. (Capsa dejflorata, Reeve, 37,X). IX.

Family ROCELLARIIDAE.
362. Rocellaria gigantea, Deshayes (Gastrochaena gigantea, Sowerby, 190,a, XX). IX.
363. Rocellaria mytiloides, Lamarck (Gastrochaena mytiloides, Sowerby, 37,XX). XII.

Family MESODESMATIDAE.
364. Paphia trigona, Deshayes (MJesodesma trigona, Reeve, 37, VIII). IX.
365. Paphia glabrata, Gmelin (Mesodesma glabrata, Reeve, 37, VIII). IX.

Family LUCINIDAE.
366. Lucina (Codakia) divergens, Philippi, 33,}. (Lucina fibula, Reeve, 37, VI). IX.
367. Lucina (Codakia) punctata, Linn. 37, 7. IX.

368. Lucina (Divaricella) dalliana, Vanatta var. 42. IX.

Three valves from the Maldives are a trifle more globose than 8. African specimens, the
divaricating lines are rather less numerous, and the smooth band, where they meet, is not quite
so evident. In typical and especially in young examples the posterior straight dorsal margin forms
a rounded angle where it meets the posterior curved outline. This angle is less apparent in the
Hulule specimens.

The very fine crenulation within the margin of the valves is similar in both forms.

This species has been erroneously quoted by Mr Sowerby as JL. quadrisuleata, d’Orbigny,
(Marine Shells S. Africa, Appendix, p. 26). Of this I speak with certainty as the shells so-named
by him are now in the British Museum.

369. Cryptodon globosus, Forskal (Lucina globosa, Pfeiffer, 32). IX.

Family TELLINIDAE.
370. Tellina elegans, Gray, 40,0, XVJ/. IX.

371. Tellina crucigera, Lamarck, var. wmoena, Deshayes, 37, XVZ/. IX.

MARINE MOLLUSCA. 627

372. Tellina rastellum, Hanley, 19. XV.
373. Tellina dispar, Conrad, 40,),/. IX.

Two specimens, one white suffused with pale yellow, the other exhibiting a broad rosy ray
posteriorly and a fainter one in front.

374. Tellina rhomboides, Quoy and Gaimard, 19,a. XV.
375. Tellina robusta, Hanley, 19,a. X, XIV, XV.

376. Tellina rugosa, Born, 40,a, XVJJ. IX.

377. Tellina scobinata, Linn. 40,a,XVJJ. IX.

378. Tellina carnicolor, Hanley, 40,6, Z (Tellina corbis, Sowerby, 40,6, 7. Tellina strangei,
Deshayes, 12,c.) III, X.

The types of 7. corbis and 7’. strangei being in the British Museum collection I have carefully
compared them with Hanley’s species and I fail to discern even varietal differences. The “more
equilateral” form in 7’. corbis spoken of by Sowerby is so very slight as to be scarcely appreciable
and may be regarded merely as an individual variation. Tellina tessellata, Deshayes from Moreton
Bay (=Z. tenwistriata, Sowerby, Con. Icon. fig. 128) is also closely allied but is somewhat differ-
ently sculptured. 7’. tenwistriata is described as “orange” in colour, but the figure is purplish
brown in the copy of the work at hand. In reality it is. of a pale fleshy tint and orange along
the dorsal margins, with some opaque white markings radiating from the umbones.

379. Tellina verrucosa, Hanley, 40,0, YV/J/. IV.

Family SCROBICULARITDAE.
380. Semele jukesi, A. Adams (Amphidesma jukesii, Reeve, 37, VZ//). VIII.

Family CUSPIDARITDAE,

381. Cuspidaria elegans, Hinds (Weaera elegans, Smith, 39,a; Neaera moluccana, Adams
and Reeve, 3). X.

LITERATURE.

1. Apams, A. (a) Proc. Zool. Soc., 1851, pp. 267 et seg.; (b) Proc. Zool. Soc. 1853,
pp. 69—74, 94—99, 116—119; (c) Thes. Conch., Vol. 11.

2. Apams, H. Proc. Zool. Soc., 1868, pp. 12—17, 288—294.
. Apams and REEVE. Voyage “Samarang.”

. AwneGas. Proc. Zool. Soc., 1872, pp. 610 et seq.

- Brecx. MSS.

3

4

5. Barrp. Brenchley’s Cruise of H.M.S. “ Curacoa.”
6

7. CANEFRI. Ann. Soc. Mal. Belaique, Vol. xv. p. 50.
8

. CHeEeNnv. Man. Conch.

10.
11.
12.

EDGAR A. SMITH.
CHEMNITZ. Conch. Cab., Vol. x.
CuLEssin. Conch. Cab., Ed. 2.
Crossz. (a) Journ. de Conch., Vol. xi.; (b) Rev. et Mag. Zool., Vol. x. 1858.
DEsHAYES. (a) Maillard’s L’Ile de la Réunion, Vol. .; (b) Journ. de Conch., Vol. v1.;

(c) Proc. Zool. Soc., 1854, pp. 317 et seg.; (d) Proc. Zool. Soc., 1859, pp. 270 et seg.

13.
14,
15.
16.
17.
18.

DELESSERT. Recueil de Coquilles.

D’Orsieny. Moll. Cuba, Vol. 11.

Dunxer. Index Moll. maris Japon.

GMELIN. Syst. Nat.

GOULD. Otia Conchologica.

Gray. (a) Proc. Zool. Soc., 1834, pp. 57 et seg.; (b) (?Gray) Appendix, King’s

Australia, Vol. 1.

19.
20.
21.
22.
23.
24.
25.
26.
27.
28.

p. 150 e¢ seq.

29.
so.
31.
32.
33s.
34.
35.
36.
37.

XVII. XX.

HaANutEy. (a) Thes. Conch., Vol. oe (b) Ipsa Linn. Conch.

Hinps. (a) Voyage “Sulphur” Mollusca; (b) Thes. Conch., Vol. 1.

Issex. Malacol. mar Rosso.

JOUSSEAUME. Rev. et Mag. de Zool., Vol. xxi.

KIENER. Cog. viv.

KoBELT. Conch. Cab., Ed. 2.

Krauss. Sudafr. Moll.

Martens. Mobius’ Meeresfauna Mauritius, Mollusken.

MeELvitu. (a) Mem. Manchester Soc., Vol. xut.; (b) Journ. Conch., Vol. x.

MELVILL and STANDEN. (a) Journ. Conch., Vol. vuil.; (b) Journ. Linn. Soc., Vol. XXVI.

MELVILLE and Sykes. Proc. Malac. Soc., Vol. 111.

Morcu. Cat. Conch. Yoldi.

PEASE. (a) Amer. Jour. Conch., Vol. 111.; (b) Proc. Zool. Soc., 1860, p. 397 et seg.
PFEIFFER. Conch. Cab., Ed. 2.

PHILipPl. (a) Conch. Cab., Ed. 2; (6) Abbild., Vol. 11.

Pruspry. Man. Conch., Vols. XI. XII. XIV. XV. XVII.

Quoy et GaimaRrD. Voyage “ Astrolabe.”

R&éciuz. Journ. de Conch., 1850.

REEVE. Oonch. Jcon., Vols. I. If. I. IV. V. Vi. Vill. IX. X. XI. XII. XII, XIV. Xv,

MARINE MOLLUSCA. 629

88. Romer. Novitat. Conch., Monogr. Venus, Vol. 1.

89. SmiTH. (a) “Challenger” Lamellibranchiata; (b) Proc. Malac. Soc., Vol. 1.; (c) Proc.
Malac. Soc., Vol. 111.; (d) Proc. Malac. Soc., Vol. v.; (e) Journ. Linn. Soc., Vol. x11.; (f) Proc.
Zool. Soc., 1878, pp. 804 et seg.; (g) “Alert” Report.

40. SowerBy. (a) Conch. Icon., Vols. XV. XVII. XVIII. XIX. Xx.; (b) Thes. Conch., Vols.
I. Il. I. Iv. v.; (c) “Zankerville” Cat., Appendix; (d) Proc. Malac. Soc. Vol. 1.; (e) Proc.
Malac. Soc., Vol. tv.

41. Tryon. Man. Conch., Vols. I. IV. VI. VII. IX.
42. Vanatta. Proc. Acad. Nat. Sci. Philad., Vol. Lm.
438. Weinkavurr. Kiister’s Conch. Cab., Ed. 2.

EXPLANATION OF PLATES.

PLATE XXXV.

Fig. 1. Cryptophthalmus minikoiensis, dorsal view.

Fig. 2. 5 3 shell.
Figs. 3, 4. Marginella pictwrata, var.

Fig. 5. Mitra (Costellaria) angustissima, n. sp.
Fig. 6. Massa maldivensis, n. sp.

Fig. 7. o6 79 operculum.

Fig. 8 3 5 var.

Fig. 9 5, mulukuensis, n. sp.

Fig: 10: 5; PA operculum.

Fig. 11. ,, subtranslucida, n. sp.

Fig. 12. ,, = disparilis, n. sp.

Fig. 13. Murex (Ocinebra) submissus, n. sp.
Fig. 14. Sistrum iostoma, A. Adams.

Fig. 15. Triforis lilaceocinctus, n. sp.

Fig. 16. . excellens, n. sp.

Fig. 17. 5 % sculpture magnified.
Fig. 18 » gracilior, n. sp.

Fig. 19. £ 33 sculpture magnified.
Fig. 20. » pura, 0. sp.

Fig. 21 43 », sculpture magnified.

630

. 22,
. 23,
. 24,
25,

ig. 27.

1,

EDGAR A. SMITH.
Natica euzona, operculum.
Scalaria (Cirsotrema) texta, n. sp.
Liotia squamicostata, n. sp.
26. Thalotia maldivensis, n. sp.

Perrinia angulifera, A. Adams.

PLATE XXXVI.

2. LHthalia rhodomphala, un. sp.

Ischnochiton hululensis, n. sp.

” © anterior valve.
” ” median ”

” ” posterior ,,

- maldivensis, n. sp.

” 7 anterior valve.
” ” median 9
” » posterior ,,

-p Jeliduensis, n. sp.

” 6 anterior valve.
” ” median _,,

” of posterior ,,

Pecten coopert, n. sp. left valve.

Fr 3 sculpture magnified,
5 3 right valve.
i: 95 sculpture magnified.

» maldivensis, n. sp. left valve.
5 5 right valve.
Janira gardineri, n. sp. left valve.
cf rf right valve,
Amussium pauciliratum, n. sp. exterior.
” 03 interior.
Cardium levisulcatum, n. sp. right valve.

- A dorsal view.

Plate XXXV.

phy Maldives

Fauna and Geogra)

E. Wilson, Cambridge.

Edgar Smith - Mollusca.

Plate XXXVI.

sa

LiQive

Vin}
Id

auns and Geography

Fa

pea EN

‘

bridge.

Wilson Cam!

E

Edgar Smith - Mollusca.

THE ENTEROPNEUSTA.

By R. C. Punnett, M.A., Fellow of Gonville and Caius College, and Demonstrator
of Comparative Anatomy in the University of Cambridge.

(With Plates XXXVIIL—XLVI. and Text-figs, 120, 121.)

CONTENTS.

Introduction, p. 631. Systematic account, p. 631. Some anatomical points, (1) the pygochord,
p. 659, (2) the lateral septa, p. 659, (3) the branchiae, p. 660, The formation of the gonads,
p. 660. Variation in Ptychodera flava, p. 662. Metamerism in the Enteropneusta, p. 669.

INTRODUCTION.

THE collection of Enteropneusta brought together by Mr Stanley Gardiner is the most
extensive hitherto made in any single locality, not only in the variety of forms, but also in
point of numbers. To Mr Gardiner’s recognition of the importance of collecting considerable
numbers and to his perseverance and zeal I owe this opportunity of making some
observations on the variation shewn by the genus Ptychodera, a field hitherto unexplored.
The material is on the whole excellently preserved, and I would here express my thanks
to Mr Gardiner for entrusting to me so interesting a collection, I am also greatly indebted
to Mr Forster Cooper for the coloured sketches from life which form the first plate to this
paper. With the account of the Laccadive and Maldive forms I have incorporated a descrip-
tion of an interesting new form kindly placed in my hands by Mr Crossland, who collected
it in Zanzibar. Many regenerating specimens occurred in Mr Gardiner’s collection. These
I hope to make the subject of a subsequent paper.

SYSTEMATIC ACCOUNT:
Fam. 1. Harrimaniidae.
(Not represented. )

Fam. 2. Glandicipitidae.
PAGE

Spengelia porosa, WILLEY, 1898 ¢ : : : : 5 : 6 . 633
Sp. maldivensis, n. sp.
Collar relatively very short with cornua of nuchal skeleton extending to
extreme hind end. Dorsal septum throughout collar. Medium proboscis septum
very short. Dermal pits very small. No median gonads in post-branchial region. 635

1 The classification adopted is that recently revised by Spengel.

81—2

632 R. C. PUNNETT.

Willeyia bisulcata, nov. gen., n. sp.

Collar and proboscis both very long. Branchial part of cesophagus small
compared with ventral portion. No dermal pits, synapticula or median gonads .

Fam. 3. Ptychoderidae.
Balanoglossus carnosus, WILLEY, 1899 : ; : ; 5 A 5
B. parvulus, n. sp.

Small form with very short branchial region. Genital pleurae small and
widely separate at collar. Gill pouches without ventral caecum. No accessory
gonads. Collar cord with continuous lumen . .

Ptychodera flava, Escuscuoutz, 1825.
Var. parva, nov. var.

Small form. Proboscis cavity large and with longitudinal muscles gathered
into distinct separate bundles. Both proboscis pores functional. Number of
dorsal roots 2—8, Branchial region short, i.e. less than 2 x collar length.
Collar cord with continuous lumen. Nuchal cornua well developed

Var. laccadivensis, nov. var.

Large form. Proboscis cavity large and with longitudinal muscles gathered
into distinct separate bundles. Number of dorsal roots 3—6. SBranchial region
on the average more than 2 x collar length. Collar cord with continuous lumen.
Nuchal cornua well developed

Var. maldivensis, nov. var.

Moderately large form. Proboscis cavity fairly large with longitudinal
muscles gathered into distinct separate bundles. Dorsal roots only 1—2.
Branchial region averages less than 2x collar length. Collar cord with con-
tinuous lumen. Nuchal cornua short

Var. saxicola, nov. var.

Medium sized form. Proboscis cavity fairly large with longitudinal muscles
gathered into distinct separate bundles. Dorsal roots 2—4. Branchial region
very short, averaging only slightly more than collar length. Lumen of collar
cord with tendency to obliteration. Nuchal cornua short

Var. gracilis, nov. var.

Small and rather slender form. Proboscis cavity fairly large with longi-
tudinal muscles gathered into distinct separate radial bundles. Dorsal roots
2—3. Branchial region short, averaging about half as long again as collar length.
Collar cord with continuous lumen. Nuchal cornua rather short .

Var, muscula, nov. var.

Small, rather stoutly built form. Proboscis cavity much reduced and longi-
tudinal muscles not gathered into distinct separate radial bundles. Branchial
region fairly long, averaging more than 2x collar length. Lumen of collar cord
with marked tendency to obliteration. Nuchal cornua rather short

PAGE

637

640

640

644

646

648

650

651

653

THE ENTEROPNEUSTA. 633

PAGE
Var. cooperi, nov. var.
Very small form with short proboscis and collar, Proboscis cavity small
and longitudinal muscles not gathered into distinct separate radial bundles.
Branchial region rather more than 2x collar length. Lumen of collar cord
with tendency to obliteration. Nuchal cornua very long, overlapping branchial
region. : : . : 5 : : E : 5 : : : 00D

Pt. viridis, n. sp.
Medium sized form. Proboscis markedly longer than collar. Prevailing
colour in life green. Proboscis cavity very small and longitudinal muscles not

gathered into distinct separate bundles. Branchial region very short. Lumen
of collar cord completely obliterated . ‘ 4 : 6 : ; . 5 ate

Pt. asymmetrica, n. sp.
Rather small form. Proboscis cavity small and longitudinal muscles not
gathered into distinct separate radial bundles. Lumen of collar cord almost
obliterated. Gonads asymmetrical, on left side only. ; . : c 2) 607

Spengelia porosa, WILLEY, 1898 (Pl. XXXVIII. fig. 13; Pl. XLI. figs. 5, 7; Pl. XLII.
fig. 11).

Locauiry, Etc. This species was found in three localities, viz. :

(a) Goifurfehendu Atoll, Goidu, where it was obtained from the sandy shore close
to the east of the island in the sand below the weed and under some square slabs of
beach sandstone in company with Pt. flava, var. muscula. (All this material was much
macerated.)

(b) S. Mahlos Atoll, Mamaduwari, where it was very common under the beach sand-
stone masses to the north of the island.

(c) Male Atoll, Hulule. A single specimen was obtained from a hole in a coral
mass dredged from 7 fathoms in the lagoon of Hulule Atoll. (Anterior end only.)

EXTERNAL FEATURES. In Willey’s specimen the proboscis was of a rich yellow colour,
the collar bright orange and the body generally of a dull yellow colour. No observations
on the colour of the Maldive specimens have reached me except of the single one from
Hulule. Mr Stanley Gardiner noted that the colour here was uniformly pale lemon. Much
stress however cannot be laid on the difference in colour from Willey’s specimen as this
Hulule individual was represented only by the pre-genital part of the animal and was a
rather small specimen.

The general appearance of the Maldive forms agrees closely with the figure given by
Willey ?99, Pl. XXVII. fig. 8). The size also is about the same though the length of
the proboscis as compared with the collar is on the whole slightly greater than in Willey’s
specimen. The relative length of the branchial region to the collar is on the average about
6, as in Willey’s animal. In the single entire specimen in the collection the total length
was 215mm. The branchial region is fairly constant in its length relative to the collar in
individuals of different sizes. The number of gill slits increases markedly with the size of
the animal. These facts are brought out in the following table of measurements :—

634 R. C. PUNNETT.

No. of Specimen.

(1) (2) (3) (4) (5) (6) (7) (8)

Length of; collar’... .:...c0c--++<. 3 3 3 2°5 2°5 2°5 15 —
5 yy PYODOSCIS .....se.00ee 55 6 5 5 45 4:5 4 _
i », branchial region ... 20 17 16 17 175 17 10°5 9
Motal@lenp thy .ceseecsseceseaseances — 215 — — — — — —

Branchial length
Collar length

No of illWslitsicrssc-cscdeeseens 116 — 110 104 102 88 74 58

veconcorcoar eeeeee 66 56 53 68 7 6:8 a —

The ‘specimens are given in order of size, (1) being the largest, (8) the smallest.

In his account Willey states that external liver saccules are not present in the allied
species S. alba (99, p. 277), “although, when fresh, the annular ridges were found to be
turgid and to present the appearance of rudimentary saccules, an appearance which was
almost entirely lost after preservation.” He was unable to state anything with regard to
this region in Sp. porosa as his only well-preserved specimen lacked the post-genital portion
of the body. External liver saccules are nevertheless present in Sp. porosa, though they are
not very prominent (Pl. XXXIX. fig. 13). That they project somewhat from the dorsal
surface may be more readily seen in transverse sections (see Pl. XLI. fig. 5). They are
also characterized by their green colour,

\
Re -

l
ye

sg mm

Fic. 120. Schematic reconstruction of anterior end of Sp. porosa, x 10. The dotted line in front of the stomochord
indicates the forward extension of the pericardial auricles. For lettering see p. 676.

INTERNAL STRUCTURE. To the full and careful account given by Willey I have little
to add, and that chiefly in regard to the post-genital region which, as has already been
mentioned, was absent in his specimen.

Proboscis. In Willey’s specimen the ventral recurrent vessel was independent of the
ventral septum. In the three specimens which I have examined it runs up to the central
complex in the anterior free edge of the ventral septum (Text-fig. 120, vrv.).

Willey states that the median muscular system extends over about 4+ of the length
of the proboscis. It is not quite so long in the Maldive forms and may be exceedingly
short (Text-fig. 120, mdv.), not extending quite so far forwards as the pericardial auricles.

THE ENTEROPNEUSTA. 635

The vermiform process is usually well-marked and coextensive with the dorso-ventral
muscles. In the instance where these muscles were reduced the vermiform process had
altogether disappeared.

Collar. Willey states that “there is no dorsal septum in the collar, except the fold
of basement-membrane associated with the first vestigial root, which is probably to be
regarded as a vestige of the dorsal septum. On the other hand the ventral septum has
an unusual forward extension, commencing a short distance behind the region of bifurcation
of the nuchal skeleton.” In the Maldive forms the dorsal septum is present though only
at the hind end of the collar (Text-fig. 120, ds.), whilst no ventral septum is to be found.

A vestigial root was found in one of the three specimens examined. It was, however,
very small and did not reach to the epidermis (Text-fig. 120, r.). Both anterior and posterior
neuropores are very deep, so that only about 4 of the collar cord is devoid of a continuous
lumen. This portion contained traces of a cavity.

In his specimen Willey found that the cornua of the nuchal skeleton extended to the
extreme posterior end of the collar. This may be the condition in the Maldive specimens,
whilst on the other hand the hindermost quarter of the collar may be without cornua.

The collar canals open into the first branchial pouch and I have been unable to find
the truncal canals described by Willey.

Trunk. The branchial and genital regions agree closely with Willey’s account.

In the hepatic region there is no internal circular musculature. The longitudinal muscles
are gathered into three bands on either side, viz. a dorsal, a ventral, and a lateral one
(Pl. XLI. fig. 5). Consequently there are in this region two longitudinal areas where no
muscle fibres intervene between the intestinal wall and the epidermis.

In the intestinal. region there is a longitudinal dorsal groove on either side which
shews slight depressions at intervals (Pl. XLI. fig. 7). The circular muscles form a complete
ring round the body here but the longitudinal muscles are wanting beneath the grooves.
The intestine itself is small and laterally compressed, the body cavity being occupied by
strands of connective tissue.

Although there are several small points of difference between the Maldivan forms and
Willey’s specimen I have thought it best, in view of the considerable variations shewn on
these very points, not to separate these forms from Willey’s. It is possible that when more
material is forthcoming the differences may be found to be sufficient to separate them as
distinct varieties.

Spengelia maldivensis, n. sp. (Pl. XLI. figs. 6, 8; Pl. XLII. fig. 20).

Locatiry, ETC. Hulule. From beach sand-rock. A single specimen only was obtained
and this lacked the posterior part of the body, which was broken away in the genital region.

EXTERNAL FEATURES. A small form and the colour after preservation is quite white.
The total length of the fragment obtained was 21mm. Of this the proboscis measured
4mm., the collar 1‘8mm. (with a width of 3mm.), whilst the length of the branchial region
was 6mm. Characteristic is the shortness of the collar, the length of which is not much
more than half its width. The width of the genital region did not exceed 3 mm.

636 R. C. PUNNETT.

INTERNAL STRUCTURE.

Proboscis. The outer circular muscle layer is very delicate. The proboscis contains a
central cavity which is not divided into right and left portions in front of the central
complex. The long vermiform process of the stomochord runs forwards dorsal to the cavity
(Pl. XLI. fig. 8, v.). It is somewhat fragmented in places. Where it joins the stomochord
proper appears the ventral septum separating the ventral coelom into two portions, which
do not subsequently jo with one another. A dorsal septum also occurs in front of the
junction of the pericardium with the basement membrane below the epidermis. The lumen
of the stomochord is well marked in some places though obliterated in others. The lumen
of the lateral diverticula is only patent at the extremity of each. The chondroid tissue is
well developed. The nuchal skeleton possesses a well-marked keel.

The left proboscis pore alone is present. The right perihaemal cavity reaches forwards
considerably further than that of the left side (Pl. XLI. fig. 6, ph.).

The pericardial auricles are very short.

Collar. The outer longitudinal muscles are poorly developed and there is no circular
layer. The collar is without a cavity. The dorsal septum extends through the length of
the collar, whilst the ventral one is only found complete at the posterior end.

The nerve cord is practically solid throughout, only very faint traces of a lumen being
discoverable here and there. There is no sign of any dorsal root.

The cornua of the nuchal skeleton are long and reach backwards to the extreme hind
end of the collar (Pl. XLI. fig. 8, cn.) and to some extent overlap the branchial region.

The collar pores open into the first gill pouch, which possesses a small diverticulum
reaching forwards for a short distance ventral to the collar canal (Pl. XL. fig. 8, p.1). This
is doubtless the structure referred to by Willey (99, p. 280) as the truncal canal. In
the present species it possesses no trace of an opening into the perihaemal space, and the
diverticulum differs in nowise histologically from the rest of the gill pouch, thus lending
no support to the morphological importance with which Willey is inclined to regard them.

Trunk. The circular musculature is, as usual in this genus, found inside the longitudinal
layer. In the branchial region the ventral portion of the cesophagus is almost as large as
the branchial division, the proportions being similar to those in Sp. porosa as figured by
Willey ’99, Pl. XXXI. fig. 45). About nine synapticula are present on each gill bar. A
distinct but small post-branchial groove is present (Pl. XLII. fig. 20). The grooves into
which the gill pouches open are produced backwards past the branchial region. They do
not however exhibit the peculiar depressions found in Sp. porosa and Sp. alba which Willey
has termed dermal pits. Too much stress must not be laid on their absence, however, as
the gonads are somewhat immature and it is possible that with their increase in size small
dermal pits might arise. There is little doubt that they would never reach the same
development as in the above-named forms, a circumstance probably correlated with the smaller
size of the species under consideration.

In the branchial region both median and lateral gonads are present. In the genital
region however the lateral gonads alone are present. The median gonads are always very
small and are not nearly so numerous as the lateral ones.

Though the above species has been created on the strength of a single imperfect
specimen yet in its short collar, its undivided proboscis cavity, the reduction or absence of

THE ENTEROPNEUSTA. 637

its dermal pits and median gonads, it exhibits structural differences sufficient to separate
it at once from Sp. porosa and Sp. alba, the only two species of this genus hitherto described.

Willeyia bisulcata, nov. gen., n. sp. (Pl. XXXVIII. figs. 14, 23; Pl. XXXIX. fig. 29;
Pl. XLI. figs. 9, 10; Pl. XLII. figs. 12—16).

Locauity, ETc. This species was fairly abundant at Zanzibar, whence portions of two
specimens were brought back by Mr Crossland. They were obtained by digging in mud
on which Zostera was growing at low-water mark.

EXTERNAL FEATURES. To judge from the fragments the length of this species must in
the entire state be considerable. The proboscis is about double the length of the collar
and is marked both dorsally and ventrally by a deep median groove (Pl. XXXVIII. fig. 14).
Its width is somewhat greater than its dorso-ventral depth. The collar is long and is
somewhat broader behind than in front. Near its posterior end there is a well-marked
circular groove. In one of the two specimens available, which was somewhat larger than
the other, the length of the proboscis was 14 mm., that of the collar 7mm. The average
breadth of both was about 4mm. The branchial region and trunk have each a breadth
somewhat less. In the specimen above alluded to some 32 mm. of the branchial region
were still attached to the collar. No external liver saccules were visible on any of the
fragments. Probably they are not present. No portion of the caudal region was preserved.
With the exception of the proboscis, which is somewhat dorso-ventrally flattened, the whole
body of the worm is more or less cylindrical. When living the animal was of a uniform
cream colour.

Proboscis. The epithelium is high though almost destitute of glands. It shews a very
distinct layer of elongated nuclei about its middle and numerous small rounded nuclei in its
deeper portion. The layer of nerve fibrils is very distinct and well developed throughout
this region and rests on a fine but clearly marked basement-membrane. The circular muscle
layer is strong, being nearly half as thick again as the nervous layer. The longitudinal
muscles are well-developed, filling the greater portion of the proboscis (Pl. XLII. fig. 12),
The nervous layer is thickened in the grooves.

The proboscis coelom consists of a right and a left cavity which are entirely separate
throughout. They are lined by a well-marked epithelium which is considerably thickened
near the middle line (Pl. XLII. fig. 12 and Pl. XXXIX. fig. 29). The two divisions of the
coelom are surrounded by a common layer of circular muscles (Pl. XLII. fig. 12). The left
division of the proboscis coelom communicates with the exterior by a large pore (Pl. XLI.
fig. 9). The right division ends blindly. No dorsal septum is present but there is a well-
marked ventral septum, at the upper edge of which runs the ventral recurrent vessel of
the proboscis (Pl. XLII. figs. 11 and 12, and Pl. XXXIX. fig. 29, wrv.). This septum
springs from almost the tip of the proboscis and does not become complete until shortly
behind where the pericardium arises. Here the ventral vessel becomes connected with the
vessels of the glomerulus. On either side of the middle line are found dorso-ventral
muscles, the dorsal ones being more marked than the ventral. Both converge towards the
vermiform process of the stomochord where this structure is found (Pl. XXXIX. fig. 29).

The stomochord exhibits a well-marked lumen throughout, both in its central portion
and in the lateral diverticula. Anteriorly it is produced forwards as a vermiform process
which is about 2 of the length of the rest of the stomochord (Pl. XLII. fig. 10).

G. IL 82

638 R. C. PUNNETT.

The nuchal skeleton is strongly developed, occupying a large portion of a transverse
section in the region of the proboscis stalk (Pl. XLI. fig. 9). The keel is strongly developed
but much blunted, its side to side diameter being greater than the dorso-ventral one.

The pericardium is provided with two small “auricles” reaching forward about ~; mm.
from the main structure on either side.

Collar. The epithelium of this region is high and fairly well provided with gland-cells.
The posterior part of the collar on either side of the circular furrow is characterized by a
more highly glandular zone. In the preserved specimens this region is somewhat paler in
colour than the rest of the collar. The nerve fibre layer is distinct, whilst, as in the
proboscis, the basement-membrane is thin. There is no layer of circular muscles outside the
longitudinal musculature.

The collar coelom is entirely filled by longitudinal muscle fibres and connective tissue,
with the exception of a small space round the opening of each of the collar funnels. The
dorsal septum is complete throughout the entire length of the collar. The ventral septum
is lacking only at the extreme anterior end (Pl. XLI. fig. 10).

The median nuchal skeleton is long whilst its cornua are exceedingly short (Pl. XLI. fig. 10),
a feature in which this genus differs markedly from the remaining members of the family.
The cornua embrace nearly ? of the total circumference of the cesophagus.

The collar cord exhibits a well-marked anterior and a smaller posterior neuropore. The
anterior neuropore especially reaches some way down into the collar (Pl. XLI. fig. 10).
Towards its hinder limit it becomes somewhat separated off from the nerve cord on the
dorsal surface of the latter. It is lmed with a definite epithelium somewhat glandular and
richly provided with cilia. The nerve cord itself is quite solid and is almost entirely sur-
rounded by a layer of nerve fibres (Pl. XXXVIII. fig. 23). In its hinder portion it shews a
well-marked ridge projecting up towards the dorsal septum. This ridge is filled with cells
containing a yellowish-brown pigment. These cells reach a little way into the nerve cord
and similar cells are also to be found in the perihaemal spaces (PI. XXXVIII. fig. 23).
Pigment has previously been described by Spengel (98, p. 134 and Taf. 7, figs. 2, 12, 14)
in the collar cord of Balanoglossus apertus. Without going so far as Willey (99, p. 316)
in claiming that the dorsal roots are homologues of the vertebrate pineal eyes it is yet
conceivable that these aggregations of pigment may serve for the perception of light rays.
Besides this small dorsal ridge there occurs no representative of dorsal roots in W. bisulcata.

From the anterior part of the collar cord passes off on either side a well-marked nerve
to the layer of nerve fibrils beneath the cesophageal epithelium. These I have termed the
cesophageal nerves (Pl. XLII. fig. 16, on.).

With regard to the histology of the collar cord it may be stated that giant cells, such
as Spengel (98, p. 609) has described for several species of Enteropneusts, are absent.
Certain of the ganglion cells near the dorsal surface attain a somewhat larger size, but
this does not exceed 20—25 yw. Faint vestiges of the remains of cavities in the cord are
visible here and there under a high power. The pigment referred to above is found along
the whole length of the cord.

The perihaemal spaces are well developed and ventrally contain transverse muscle fibres.
They do not come into contact with one another until just in front of the point where
the cornua of the nuchal skeleton diverge. Anterior to this they are widely separated by
the stomochord.

THE ENTEROPNEUSTA. 639

The right and left peripharyngeal spaces are separate except at the hind end of the
collar, where they meet ventrally. As they reach forwards their ventral borders slope obliquely
and gradually upwards until the two borders merge at a spot dorsal to the cesophagus,
where the circular muscles which the spaces contain are inserted into the nuchal skeleton.
Where the peripharyngeal spaces are deficient ventrally the wsophagus is still provided with
a layer of circular muscles probably derived from the longitudinal musculature.

Branchial region. One of the chief characteristics of the present species as compared
with the other members of its family lies in the fact that the branchial portion of the
esophagus is of considerably less extent than the remainder (Pl. XLII. fig. 13). This results
in the septal and tongue bars being relatively short. The musculature is in this region
well-developed. Beneath the powerful longitudinal layer is a sheath of circular fibres extend-
ing up to the lower ends of the gill pouches. Noteworthy is the position of the ventral
nerve cord which projects so far inwards that it almost abuts against the circular muscles.
Of the posterior branchial region it is not possible to give any account, since I have been
unable to identify it among the fragments which have come to me. Synapticula are absent.
The first three gill pouches are dorsally confluent, forming a chamber with a single external
pore into which opens the collar pore. No trace of truncal pores is to be found resembling
those described by Willey (99, pp. 273, 280) for Spengelia. The branchial region is
probably devoid of gonads, or if these occur here it is only in its most posterior portion
that they are to be found. In the specimen with the greater attached length of branchial

region, this was about 44 times the length of the collar.

at the extreme end of this fragment.

No gonads were present even

Schizocardium

Glandiceps

Spengelia

Willeyia

Pericardial auricles very
large.

No dermal genital pits.
chial sac very small.

Peripharyngeal cavities.

Cornua of nuchal skeleton
to posterior collar re-

gion.

No median gonads.

Synapticula.
External liver saccules.
Ventral proboscis septum

long and reaching ver-
miiform process.

(Esophageal part of bran- |

Pericardial auricles very
small.

No dermal genital pits.

(Esophageal part of bran-
chial sac reduced to a
broad tract.

Cornua of nuchal skeleton
to posterior collar re-
gion.

Median gonads present or
absent.

No synapticula.
No external liver saccules.
Ventral proboscis septum

short and not to vermi-
form process.

No peripharyngeal cavities.

Pericardial auricles small.

| Dermal genital pits.

(Esophageal part of bran-
chial sac a deep groove.

Peripharyngeal cavities.

Cornua of nuchal skeleton
to posterior collar re-

gion.

Median gonads present or

absent.
Synapticula.
External liver saccules

present or absent.

Ventral proboscis septum
short and not to vermi-
form process.

Pericardial auricles small.

No dermal genital pits.

(Esophageal part of bran-
chial sac much larger
than branchial part.

| Peripharyngeal cavities.

Cornua of nuchal skeleton
not found in posterior
half of collar.

No median gonads.

No synapticula.

(?)

Ventral proboscis septum
very long but not to
vermiform process.

Genital region. The longitudinal muscles are still strongly developed (Pl. XLII. fig. 14)
but the circular layer has now disappeared. The ventral nerve cord still shews the peculiar
82—2

640 R. C. PUNNETT.

deeply sunk position which it exhibited in the branchial region. Subgenital pits such as
Willey found in Spengelia (99, p. 274) are absent. The gonads form a single row on
either side of the body. Ripe spermatozoa occurred in the specimen from which the sketch
of the genital region was taken.

As belonging to the family Glandicipitidae there have already been described three
genera, viz. Schizocardiwm, Glandiceps, and Spengelia. It cannot be said with any confidence
that Willeyia bears a closer resemblance to any one of these genera than to another.
Features which separate it from all these three are the shortness of the nuchal cornua,
the comparatively small extent of the branchial portion of the csophagus as compared with
the digestive, and possibly also the absence of gonads in the branchial region. With respect
to other features of generic importance Willeyia resembles now one genus and now another.
To which of the above genera it bears most resemblance with respect to the more salient
features of its anatomy may be gathered from the appended table, p. 639.

Balanoglossus carnosus, WILLEY, 1899 (Pl. XX XVII. fig. 3).

Locauity, ETc, The greater portion of a very large specimen was obtained at Minikoi,
at the base of a cylindrical casting in the sand flat lagoon. Fragments of the same species
were also found at Hulule in sand under a large Porites mass close to the boulder zone.

A coloured sketch was made of the Minikoi specimen by Mr Forster Cooper before
preservation. This is reproduced on Pl. XXXVII. fig. 3, and shews the natural colours of
the animal. A bright yellow line occurred mid-ventrally which was probably the nerve cord.

The dimensions of this large specimen were taken by Mr Gardiner while it was yet

alive. I have also taken them on the preserved specimen and the results of both are as
follows :—

Living Preserved
Length of proboscis beyond collar... 9 mm. 3 mm.
Breadth chit le RopOnGpaHORcooasoccaae IB} say —
gerig thmoficollarensesssaeeteseseseec tere 22, Sie
55 », branchial region ............ — 60);
: », genital pleurae............... — 204 ,,
. my Chovbootsll foJoys=nbets!6) Goopgppsqoos 440 ,, 355

When alive and undamaged the animal probably exceeded 60 cm. in length, and possessed
an average diameter of 10—12 mm. The above measurements agree fairly well with those
given by Willey (99, p. 248). There appears to be considerable variation in the relative
length of the branchial region which may be somewhat less than double the collar length,
or may be as much as four times as much. The present specimen agrees with the largest
of Willey’s specimens, which it rivals in size, in having a relatively long branchial region.

In addition to the above fairly complete specimen a number of jelly tubes were found
at the surface in castings similar to that which marked the position of the large worm.
These tubes were much distended with sand, and of a transparent straw colour with bright
yellow mid-dorsal and mid-ventral lines. The edge of the free (= anal?) end was marked
by brown-black pigment. Doubtless these tubes are the broken off caudal ends of this
species. Like Pt. flava (see p. 646) it may have developed the habit of autotomy.

Balanoglossus parvulus, n. sp. (Pl. XX XVIII. figs. 15, 18, 22; Pl. XXXIX. fig. 24;
Pl. XLI. figs. 1—4),

Locality, ETc. Represented only by a fragment comprising the anterior end and including
the proboscis, collar, branchial region, and a portion of the genital region, From Mahlos Atoll.

THE ENTEROPNEUSTA. 641

EXTERNAL FEATURES. An exceedingly small form. The proboscis is nearly 15mm. in
length and is somewhat longer than the collar (Pl. XX XVIII. fig. 18). The branchial region
is about half as long again as the collar. Width of collar and proboscis about 1°5 mm.
The width of the branchial region is not quite so great, whilst the genital region is less
than 1 mm. wide. The anterior border of the collar is rather wider than the rest of this
part of the animal and is somewhat crinkled. Posteriorly the collar shews a well-marked
circular furrow (Pl. XXXVIII. fig. 18, tf). The genital pleurae are small and are widely
separated from each other where they join the collar. The mature and swollen gonads in this
specimen cause the edges of the pleurae to assume a somewhat beaded appearance.

Proboscis. The proboscidial coelom is almost entirely filled by connective tissue and
muscle fibres. A delicate layer of longitudinal muscle fibres is found beneath the basement
membrane, but circular fibres are absent. Traces of a dorsal septum are to be found slightly
in front of the pericardial sac. The ventral septum is complete at a level shortly before
that at which the glomerulus ends; it carries the ventral glomerulus vessels. The ventral
proboscis coelom is packed with muscle fibres and connective tissue except at its extreme
posterior end where the ventral septum is lacking (Pl. XXXVIII. fig. 22, vc.). A racemose
organ is not present.

A single median proboscis pore is present, placing the left division of the dorsal proboscis
coelom in communication with the exterior (Pl. XXXVIII. fig. 22, p.p.). The right moiety of
the dorsal proboscis coelom ends blindly. Throughout its whole extent the stomochord shews
a well-marked lumen. ‘The lateral diverticula of the stomochord are small (Pl. XXXVIII.
fig. 22, s.c.). The nuchal skeleton is provided with a deep narrow keel (Pl. XLI. fig. 3, sk. 1).
The chondroid tissue in this region is exceedingly scanty.

Collar. The collar epithelium is very thick (Pl. XX XIX. fig. 24, ep.). The glandular zone
commences at the level where the cornua of the nuchal skeleton terminate and reaches to
the commencement of the collar funnels, The outer longitudinal muscle layer is well-marked,
particularly in the anterior and middle regions of the collar (Pl. XXXIX. fig. 24, el.). The
inner longitudinal layer is also strongly developed. No circular muscle fibres are present.

The collar coelom is entirely filled with muscle fibres and connective tissue except for
a very small space on either side of the dorsal septum. This last-named structure is first
found immediately behind the first dorsal root in the middle of the collar region. Traces
of a ventral septum exist but in no place is it complete. The collar funnels are of the
usual form and open into the first gill pouch.

The cornua of the nuchal skeleton extend half-way round the cesophagus. In length
they are short, being only about } of the total collar length.

The collar cord is tubular containing a single axial canal (Pl. XX XVIII. fig. 15) sur-
rounding which are the ganglion cells. The layer of dorsal ganglion cells (Pl. XXXVIII.
fiz. 15, ged.) is thin, being best marked in the region where the first and second dorsal
roots are given off. Above it is a much attenuated layer of nerve fibrils. The ventral
ganglion cells (gcv.) are much more numerous and in transverse section are seen as a V-shaped
mass shewing a bifurcation at the point of the V. The layer of nerve fibrils surrounding
them is well-developed. Separating the dorsal and ventral ganglion cells on either side is
a layer of large oval vacuolated cells—probably gland cells from which the secretion has
been ejected or dissolved out. Some deeply staining mucoid substance is to be found in
the medullary canal. Here and there occur ganglion cells rather larger than the rest,

642 R. C. PUNNETT.

but no giant cells are present such as have been figured by Spengel for B. apertus
(?98, Taf. 7, fig. 18). Except at its anterior end the collar cord is sunk in the perihaemal
spaces which embrace it on either side and which may posteriorly reach nearer to the dorsal
surface than the cord itself. The medullary canal communicates with the exterior by the
anterior and posterior neuropores.

The first two dorsal roots are large, close together, and contain nerve fibrils but are
devoid of a lumen. They spring from a common crest and reach the epidermis where their
nerve fibrils fuse with the nervous layer of the skin. Behind these are two or three small
incomplete roots with traces of a lumen but without nerve fibrils.

A well-marked peripharyngeal space surrounds the cesophagus (Pl. XXXIX. fig. 24, pph.)
reaching dorsally to where the cornua of the nuchal skeleton diverge, and ventrally somewhat
anterior to this point.

Branchial region. This region is characterized by the small genital pleurae which
are widely separate at their junction with the collar. At the base of the pleurae open
the gill pouches (Pl. XLI. fig. 1, bp.) which are not provided with ventral caeca. The branchial
and digestive portions of the cesophagus are approximately of equal size. Gonads are not
found in the anterior fourth of the branchial region. Not more than six synapticula are
present on either side of the tongue bars. The post branchial canal is-large and the posterior
incipient gill slits open into its ventral portion (Pl. XLI. fig. 2, *),

Genital region. The great bulk of this region is occupied by the gonads in the
much swollen pleurae (Pl. XLI. fig. 4). Accessory gonads are not present, but each gonad is
roughly two-branched, one division lying on either side of the lateral septum. The genital
pores are found on the edges of the pleurae at the spot where the lateral septum joins
the epidermis. The single specimen was an adult female.

Affinities. The genital pleurae of B. parvulus are relatively smaller than in any other
member of the genus and in this respect it approximates to the genus Glossobalanus. Of
the members of its own genus it bears most resemblance to B. apertus (Spengel, 798).
In both the genital pleurae are widely separate where they join the collar, and this character
serves to distinguish these two species from all the other members of the genus except
B. misakiensis (Kuwano, ’02). Both B. parvulus and B. apertus are small forms without
accessory gonads, with a tubular collar cord, and with gill pouches unprovided with a ventral
caecum. The present species however differs from B. apertus in the following features.

B. parvulus | B. apertus
Branchial region but little longer than the collar. Branchial region several times as long as collar.
Gonads open at edges of genital pleurae. Gonads open on inner surface of genital pleurae.
No circular muscles to body-wall. Feeble circular musculature.
Tongue bars with not more than 6 synapticula. | Tongue bars with about 10 synapticula.
Medullary tube open at either end. Medullary tube closed at either end.
Giant ganglion cells not found in collar cord. Giant ganglion cells present in collar cord.

In B. parvulus also the keel of the nuchal skeleton is more marked, whilst the ventral
proboscis coelom has undergone greater reduction. An idea of the relations of these two
species with the other members of the genus may be gathered from the appended table.

643

THE ENTEROPNEUSTA.

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Aypuryq spue poyouraq aeypoo
avs dTUTO[900 Arewtatg tamoaro ye yorquoo ut IeTOD X Z—T TASUOT /
a qenquy, | yf ‘etod uerpayy | “Aaosseooe ~~ ony [eaquea jNOyIIAA | you pur ]jeuIg ="yourlg | doygea«stosoqorg |jeus A1oA |***** snynawnd
(4joyem1x01ddv)
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UOIHaL [BIYOUB.q
JO sqyS09] eATyRTOYy

JO syydue] eat} RIAIT

644 R. C. PUNNETT.

Ptychodera flava, var. parva (Pl. XXXVIII. figs. 16, 20, 21; Pl) XXXIX. fig. 26;
Pl. XLII. figs. 21—28; Pl. XLIV. figs. 30—32; Pl. XLVI. fig. 48).

Locauity, etc. From Turadu, 8. Mahlos Atoll. About a dozen specimens procured, all
more or less of the same size. They were dredged from 20 fathoms on a bottom of
coarse sand.

EXTERNAL FEATURES. A small, somewhat short and slender form. The proboscis is of
about the same length as the collar, which measures 2°5—3 mm. long, and is of approximately
the same width. Behind the branchial region the body is not more than about 2mm. wide.
The liver saccules are not large and are slightly lobulated in the larger specimens. There
is no apparent differentiation in colour among them as is usual in this species. The average
length of the branchial region is about 14 times the length of the collar, it being thus
comparatively short (wide p. 673). In the single entire specimen the total length was
only 30 mm,

INTERNAL STRUCTURE.

Proboscis. A delicate layer of circular fibrils surrounds the longitudinal muscles which
are gathered together into radial bundles. A few longitudinal fibres are more closely gathered
round the central lumen of the proboscis and give it a definite outline (Pl. XLIII. figs. 21, 22).
Dorso-ventral muscles are always found dorsal to the central complex anterior to the peri-
eardium. The ventral proboscis septum is well-marked and reaches forwards almost or quite
to the tip of the stomochord. Both dorsal divisions of the dorsal proboscis coelom are in
communication with the exterior by the paired and equal proboscis pores (Pl. XLIII. fig. 26).
In one case the proboscis pores opened together almost in the mid-dorsal line (Pl. XLII.
fig. 23), a condition of great rarity in this genus and recalling those obtaimmg im Balano-
glossus apertus (Spengel, 794) and B. jamaicensis (Willey, 799, p. 293).

The racemose organ is usually well-marked but is never lobulated. The lumen of the
stomochord anterior to the lateral diverticula may be fairly distinct or almost obliterated.
The lumen of each lateral diverticulum is large and open at the end but the portion by
which it would communicate with the central lumen is usually almost or quite obliterated.

Collar. Beneath the basement-membrane is found a fine but definite layer of longi-
tudinal muscle fibres. In the anterior part of the collar they are rather better developed
and here directly enclose a delicate layer of circular muscles. No outer circular layer is
found over the greater part of the collar. The anterior free cavity of the collar coelom
is very spacious (Pl. XLIII. fig. 27,c.) extending as usual completely round the cesophagus
in this region. Inside it are found the inner longitudinal muscles of the collar and within
these again the peripharyngeal spaces with their enclosed circular muscle fibres. The dorsal
collar septum is as usual complete immediately behind the first root. In one case it was
complete just anterior to this structure. The ventral septum may be absent or present in
the extreme hind portion of the collar.

The nerve cord exhibits a complete lumen throughout, though sometimes with a faint
tendency to obliteration. When this occurs the height of the cord as seen in transverse
section is rather greater than usual and the gland cells are more numerous (cf. Pl. XX XVIII.
figs. 20 and 21). The number of the dorsal roots is either two or three. The majority
reach to the ectoderm.

The cornua of the nuchal skeleton extend over nearly a quarter of the length of the
collar, Their length is apparently very fairly constant for this variety (cf. Table 1, p. 645).

THE ENTEROPNEUSTA.

TABLE 1.

Ptychodera flava, var. parva.

645

8

v

No. of specimen (1) (2) (3) (4) (5)
IPLODOSCIS|COCLOM:......-c.ccoscss eens Spacious. Ca- as (1) as (1) as (1) as (1)
vity with dis-
tinct outline
Collamicoclomirccscccsesssesseseees Very spacious as (1) as (1) as (1) as (1)
Ventral proboscis septum ......... To tip of sto- as (1) as (1) as (1) Almost to tip
mochord of stomochord
Dorso-ventral proboscis muscles Present as (1) as (1) as (1) as (1)
Dorsal collar septum ............... Complete after as (1) as (1) Complete just as (1)
1st root before 1st root
Ventral collar septum ............ Absent as (1) Complete — at as (3) as (1)
hind end of
collar
IRACEMOSE OFFA) .c.ecsccesssessescces Small and un-| Well marked as (2) as (2) as (2)
lobulated but unlobu-
lated
Dorsal proboscis muscle decussa- | Small. No ven- as (1) as (1) as (1) as (1)
tion tral circular
ruse. fibres
WorsaliOotsenc.-s-cectereseecaseer *Rrr RRr RR Rr RR
IETOUOSCISIPOLESI c= sdecesertaedcessee *LR LR LR LR LR (opening
together)
IN@GVe;COrdy sessaeaccceeeressdesoesess Lumen com-| Lumen with} Lumen with as (1) as (1)
plete through- | fainttendency | very faint
out to occlusion tendency to
occlusion
Length of post-branchial groove e
= “16 — . ‘16
Length of branchial region 2 i Ie
fpbceed Moonen a ee 21-2 26°5 22'8 21-7 21-0
| Length of collar
|
| y | He je 5
Circumference of branchial ces. Vi 1-05 1-06 1-04 1-07
Circumference of cesoph. ces.
| PSHVME SALLI Goccessoodooabcoscenoqdan6 Not more than | Not more than as (1) as (1) as (1)

* R denotes a dorsal root reaching to the epidermis, r an incomplete root which does not extend to

the epidermis.

and places the proboscis coelom in communication with the exterior a capital letter is used.

pore is not thus functional it is denoted by a small letter.

Of the proboscis pores L represents the left and R the right one. When the pore is functional

When the

Trunk. The branchial region, as has already been seen, is in this variety relatively

short.
of the branchial region.

G. IL.

The post-branchial groove is moderately well developed and is about }
The genital folds in the region of the branchiae are large

of the length

83

646 R. C. PUNNETT.

(Pl. XLVI. fig. 48), springing from low down. The branchial division of the cesophagus is
somewhat more capacious than the ventral division of the cesophagus. The number of
synapticula on the longest gill bars is either 7 or 8.

Ptychodera flava, var. laccadivensis (Pl. XXXIX. fig. 25; Pl. XL. figs. 36, 38—40:
Pl. XLV. fig. 43; Pl. XLVI. figs. 49, 51, 57).

Locauity, ETc. Muinikoi; from lagoon close to edge of lagoon flat. It occurred in 4
feet of water at low spring tides and was very abundant. The followmg notes were made

by Mr Gardiner.

“A few points about them are of interest, viz. (a) The tail part of the body, ie.
posterior 1, is merely a bag of sand and is often broken off if the animal be held still
by the head end. Under these circumstances the sand appears to be driven back by peri-
staltic movements of the whole body-wall more rapidly than it can escape, and as a result
the posterior end simply breaks off. This I have verified by repeated observations under
water and I believe that it occurs im life. (b) In distribution they are limited to the
weed and to an area close to the edge of the flat. (c) Their natural position when alive
is within 4 inches of the surface of the sand. The head is close to the surface whilst
the body and tail are horizontal. (d) When thrown on to a sandy bottom the motion is
very feeble or nil if the body be full of sand. The posterior part is at once dehisced. A
wriggling motion of the branchial and genital regions ensues, accompanied by powerful con-
tractions of the collar and extensions of the proboscis. The proboscis and collar! are very
slowly forced into the sand and then the body follows with comparative rapidity.”

EXTERNAL FEATURES. This variety shews a great resemblance to Pt. flava, var. cale-
doniensis (Willey, 799, p. 228) in general appearance. The largest complete specimens which
reached me are about 150 mm. in length, but fragments of larger specimens point to its
attaining a somewhat greater size. The proboscis is cf about the same length as the collar
or slightly longer. The anterior liver saccules are dark, the posterior lighter in colour. The
larger liver saccules are lobulated along one edge, such lobulation being most marked in
the larger specimens. The length of the branchial and post-branchial genital regions are
dealt with later (p. 663) in connection with variation in the group. The colour of the
animal in life was bright yellow or pale lemon, thus differing somewhat from Pt. caledoniensis,
which is of a duller hue inclining to brownish.

INTERNAL STRUCTURE.

Proboscis. Just beneath the basement-membrane occur a few delicate circular fibres
surrounding the longitudinal muscles which are gathered up into radial bundles. These
bundles project into the proboscis coelom, which has no well-defined boundary (Pl. XXXIX.
fig. 25). Dorso-ventral muscles are usually to be found in the region of the central complex.
The ventral proboscis septum extends forwards as far as the tip of the stomochord. The
decussation of muscle fibres on the dorsal side of the anterior portion of the proboscis is
well-marked as a rule, and a few circular fibres may be continued from it round the inner
edge of the longitudinal muscles on the ventral surface of the proboscis cavity. The proboscis
coelom of both sides may be placed in communication with the exterior by the two proboscis
pores. It as often happens however that the communication between the proboscis coelom

1In Dolichoglossus pusillus, whose movements form the subject of a paper by Ritter (02), the proboscis appears
to play a relatively more important part.

and the end vesicle opening by the proboscis pore is lacking.

one side and sometimes on

THE ENTEROPNEUSTA.

the other.

TABLE 2.

Pt, flava, var. laccadivensis.

647

Sometimes this occurs on

(2)

No. of specimen (1) (Regenerated) (3) (4)
Proboscis CoelOM!..< 22... .ccce.-n eee Fairly large as (1) Large Spacious
Collsricoclomies...5.c..2c0e-ce-s-eeee Spacious Small but as (1) as (1)

well defined
Ventral proboscis septum ......... To tip of — as (1) as (1)
stomochord
Dorso-ventral proboscis muscles Absent _ Traces Present
Dorsal collar septum ............... Complete | (Regenera- as (1) as (1)
after 1st ting)
root
Ventral collar septum ............ Absent Complete at as (1) as (2)
hind end of
collar
Racemose OrgaM ........+..0..0ee0e- Large and | Small and_ | Well marked | Well marked.
lobulated | unlobulated | and slightly | Unlobulated
lobulated
Dorsal proboscis muscle decus- | Strong with | Feeble. No | Distinet. A | Distinct but

sation

Dorsal roots

Proboscis pores

Nerve cord

Bee e eee were nen ee eeneennee

Length of post-branchial groove
Length of branchial region

Length of cornua x 100
Length of collar

Circumference of branchial ces.

ventral circu-
lar muscles

*RRRRRR

*1R

Lumen
continuous

bo
i)
1

Circumference of cesoph, ces.

Synapticula

ventral circu-
lar muscles

Collar cord
in contact
with
epidermis
throughout

LR

Middle of
cord solid

few ventral
circular
muscles

RRRr

LR

Lumen
continuous

no ventral
circular
muscles

Rrrr

LR

Lumen
continuous

as (4)

After Ist
root. In-
complete in

places

as (1)
as (1)

Strong with
a few ventral
circular
fibres

RRr

Lr

Lumen with
strong ten-
dency to
obliteration

22°5

* See foot-note on p. 645.

as (2)

as (4)

Well marked.
No ventral
circular
muscles

RRR

IR

Lumen
continuous

on
cr

83—2

648 R. C. PUNNETT.

The racemose organ is always well-marked and in the larger specimens is usually some-
what lobulated.

Collar. The cavity in the anterior part of the collar is usually spacious though not
quite to the same extent as in Pt. parva. The dorsal septum is usually complete after
the first root. The ventral septum may be entirely absent or, equally as often, it may be
found in the hinder part of the collar. The number of proboscis roots varies from three
up to as many as six. The collar nerve cord usually has a distinct and continuous lumen
which may rarely shew a tendency to become obliterated. The length of the cornua of
the nuchal skeleton is subject to a good deal of variation. It averages a trifle less than
a quarter of the collar length.

Trunk. In the branchial region the number of synapticula varies with the size of the
animal. The branchial region averages rather more than double the length of the collar
(p. 665, Table 8). The post-branchial groove is well-developed, its length being about one-fifth
of that of the branchial region. At its anterior level the genital folds are still of con-
siderable size (Pl. XLVI. fig. 49), At the end of the post-branchial groove the folds have
become much reduced (Pl. XLVI. fig. 51). The branchial portion of the cesophagus is
somewhat smaller in section than the cesophageal portion (Pl. XLVI. fig. 57). A pygochord
is present in the caudal region.

Ptychodera flava, var. maldivensis (Pl. XX XIX. fig. 27).

Locauity, ETc. From Maradu, Addu Atoll, where it was taken from sandy holes of the
reef-flat. Fairly abundant. Two specimens also obtained from Hulule, on the east reef
underneath the stones of the boulder zone in sand.

EXTERNAL FEATURES. In general shape and appearance this variety is indistinguishable
from laccadivensis to which it is very closely allied. Moreover the coloration in life was
similar, this variety, like laccadivensis, shewing differentiation of the hepatic caeca into an
anterior dark group and a posterior larger and lighter group.

INTERNAL STRUCTURE. Exceedingly similar to that of Jaccadivensis from which variety
however the present one would seem to differ on the following points. The proboscis cavity
has a definite outline as in parva (cf. Pl. XLIII. fig. 21) and does not reach up among
the radial muscle bundles as in laccadivensis (cf. Pl. XXXIX. fig. 25).

The racemose organ is on the whole somewhat smaller.

The dorsal roots are Jess numerous, a feature in which this variety resembles cooperv.

The cornua of the nuchal skeleton are relatively shorter than in laccadivensis.

The branchial portion of the cesophagus is as large or larger than the ventral portion,
whilst in laccadivensis it is always smaller.

The branchial region is relatively shorter than in laccadivensis, averaging only 1°63 times
the collar length as against 2°26 times in the latter variety, a difference which is the more
striking when one takes into account the comparatively large number of specimens examined
(see below, p. 665, Table 8).

The post-branchial groove of maldivensis is also somewhat shorter than that of laccadivensis.

In point of size the present variety is considerably smaller than Jaccadivensis, preserved
specimens of which may be as much as 150 mm. in length, whilst the largest specimen of

THE ENTEROPNEUSTA.

649

TABLE 3.
Pt. flava, var. saxicola. var. maldivensis.
No. of specimen (1) (2) (3) (4) (1) (2)
Proboscis coelom..............-.-s0s» Spacious Not large | Fairly large as (3) Fairly well as (1)
developed
and with
definite
outline
Wollarteoelomipmercc. sccsccecoecac asec Spacious Somewhat | Spacious Spacious Very as (1)
occluded by spacious
connective
tissue
Ventral proboscis septum ......... Nearly to as (1) as (1) as (1) Nearly to | Not nearly
tip of tip of to tip of
stomochord stomochord | stomochord
Dorso-ventral proboscis muscles | Present Traces as (2) Absent Present as (1)
Dorsal collar septum ...........0... Commences | Commences | Complete as (3) Complete Complete
just anterior |°5 mm. behind) after 1st root after 2nd | after Ist root
to 1st root Ist root root
Ventral collar septum............... Complete in as (1) Present in as (1) Absent as (1)
hinder part hinder part
of collar of collar but
not complete
IRACeMOSC OLEAN <.c.-caccscessesssers Well marked | Well marked | Well marked | Distinct but || Small and as (1)
butsmall and and and slightly |not lobulated || unlobulated
unlobulated | lobulated lobulated
Dorsal proboscis muscle decus- Distinet | Strong. Also|Well marked.) Strong but || Feeble. No as (1)
sation a few ventral | A few ventral} no ventral || ventral cir-
circular cireular cireular cular fibres
fibres fibres fibres
WMorsalinoots|s.cccn.ccs. casssecosseacoss *RRrr Rr RR RRR Rr R
IRTODOSCIS| POLES) sere cascecescecease=: *Lr LR IR LR Lr IR
WNervercord! (22 esscccssesceecaseses Lumen with | Lumen with Lumen Lumen with Lumen Lumen
tendency to | marked ten-| continuous | slight ten- || continuous. | continuous
obliteration | dency to dency to Slight ten-
in middle | obliteration obliteration dency to
part obliteration
posteriorly
Length of post-branchial groove ; a ee as Eyl
Length of branchial region Fe a are 20 aS i
Length of cornua x 100 = , : = be. ;
Lenethofcollar 7077 yA 101 194 70 15°0 84
= es of branchial ces. 33 5 5 38 1-01 1:20
Circumference of cesoph. ces.
SVU UICUIA ees vececnnseeccieessésconess Not more Not more Up to 9 Not more | = =
than 5 than 6 than 3
Length of branchial region is 2
bts : 1-2 : 95 | 7
Tengen of collar 95 5 1-16 92 134 1:70

* See foot-note on p. 645.

650 R. C. PUNNETT.

maldivensis procured was only 80mm. long. Maldivensis is also a more stunted form, its
total length averaging about 18 times the collar length as against 23 times in laccadivensis.

All of which considerations point to the fact that we are here dealing with two distinct
though closely related forms.

Ptychodera flava, var. saxicola (Pl. XXXVIII. fig. 17; Pl. XXXIX. figs. 28, 30, 31;
Pl. XLIV. figs. 33—41).

Locaity, ETc. Hulule, Maldive Is. Four specimens were obtained by breaking up a
dead coral mass on the sand-flat.

EXTERNAL FEATURES. This is a relatively short stunted form. The length of the
smallest specimen when preserved was 32 mm., that of the largest 65 mm. The animal is
thick in comparison with its length (Pl. XXXVIII. fig. 17). The total length relative to
the collar as unity in the four specimens is 16:2, 12°8, 165, 163, giving an average of
rather less than 16, which is low in comparison with such forms as laccadivensis (see p. 664).
The genital pleurae are large but the liver saccules are small and not lobulated. Neither
is there any differentiation into an anterior darker and a posterior lighter group.

INTERNAL STRUCTURE.

Proboscis. The radial arrangement of the longitudinal muscles is almost as well-marked
as in laccadivensis. The proboscis cavity is fairly spacious. Dorso-ventral muscles in front
of the pericardium may or may not be present. The ventral proboscis septum reaches
nearly to the tip of the stomochord. The dorsal muscular decussation in the anterior part
of the proboscis is well-marked and a few ventral fibres may pass from it round the cavity
of the proboscis. Paired proboscis pores are present of which one or both may be in
functional communication with the dorsal coelom. The racemose organ is well-marked but
differs considerably in size among the different specimens. It may shew a certain amount
of lobulation.

Collar. The cavity of the anterior part of the collar is spacious. The dorsal collar
septum in two cases shewed the usual condition, ze. was complete after the Ist root. In
one case, however, it commenced about ‘5 mm. behind the Ist root, and in another a little
in front of it. The ventral collar septum is more marked than usual, being in three of
the four specimens complete in the hinder part of the collar.

The nerve cord may shew a lumen continuous throughout or there may be a greater
or less amount of occlusion. The number of dorsal roots varies from 2—4, In one specimen
a somewhat remarkable dorsal root occurred. It was of considerably larger size than usual
(Pl. XXXIX. fig. 28) and was obviously formed by two distinct portions, viz. an outgrowth
from the collar cord with a cap of yellowish brown pigment, and an epidermal ingrowth
at the end of which was a large mass of mucus. The two portions of the root fused more
or less with one another. It is possible to regard such a structure as possessing some
sensory significance, and to suggest that the mucous portion of the epidermal ingrowth may
act as a crude lens in focussing rays of light on the pigmented portion of dorsal outgrowth
from the collar cord.

The cornua of the nuchal skeleton vary greatly in length but are on the whole un-
usually short.

Trunk. One of the most conspicuous features of this variety is the small size of the
branchial region. Not only is the branchial portion of the cesophagus very small compared

THE ENTEROPNEUSTA. 651

with the ventral portion (Pl. XLIV. fig. 38), but the length of the branchial region compared
with that of the collar is exceedingly short. In none of the specimens is it more than a
quarter as long again as the collar, whilst in two cases it is even shorter than the latter
structure.

The post-branchial groove however is well-developed, being on the average about one-third
of the length of the branchial region.

The number of synapticula is very variable. As many as nine or as few as three may
be present.

The genital folds are large (Pl. XLIV. figs. 38, 39) in the branchial region and also
in the region of the post-branchial groove (Pl. XLIV. fig. 41). Just behind this region
they rapidly diminish in size (Pl. XLIV. fig. 40). In all four specimens the gonads are
quite immature. Points of interest in connection with them are dealt with later (p. 661).

An account of the pygochord is given on p, 659.

Ptychodera flava, var. gracilis (Pl. XX XVII. fig. 5; Pl. XX XVIII. figs. 10,19; Pl. XLV.
figs. 42, 44; Pl. XLVI. figs. 50, 53—55).

Locatity, ETc. Hulule, Maldive Is. From boulder zone under stones in sand. Also
from sand at Minikoi in company with Pt. flava, var. laccadivensis.

With reference to the specimens from Hulule Mr Stanley Gardiner has made the
following note:—“They are found sparingly under the stones of the boulder zone in the
sand. A couple of specimens or more can almost invariably be obtained wherever a large
grey Actinian lives, the latter seeming to protect the Enteropneusts. Polychaets, Sipunculids,
and Holothurians are not generally found in the same patch of sand.”

EXTERNAL FEATURES. This is a small variety, the longest specimen of which measured
68 mm. after preservation, the shortest 32mm. (see Table 12, p. 673). It is somewhat slenderly
built though the total length in comparison with that of the collar averages 196. The
posterior portion of the animal’s body is usually very slender (ef. Pl. XXXVII. fig. 5) though
this is not so much the case in larger specimens. The liver saccules shew an anterior dark
group and a more extensive posterior lighter group. The saccules are not large (Pl. XXXIX.
fig. 19) and are as a rule not at all or but slightly lobulated except in the case of large
specimens (Pl. XXXIX. fig. 10, *). In the specimen from which the coloured sketch
(Pl. XXXVII. fig. 5) was made all the liver saccules appear dark. It is possible that it
is only after preservation that the darker anterior group can be separated from the rest.

INTERNAL STRUCTURE.

Proboscis. The musculature is rather feebly developed, the longitudinal fibres being
gathered into widely separated radial bundles as in laccadivensis (cf. Pl. XXXIX. fig. 25).
The dorsal muscular decussation may be feebly developed or fairly strong. Dorso-ventral
proboscis muscles are present anterior to the pericardium. The ventral proboscis septum is
well-developed, reaching almost to the tip of the stomochord. Of the paired proboscis pores
only one is functional, placing the dorsal coelom in communication with the exterior. This
may be either on the right or left side. The other pore is generally smaller and in one
case had disappeared altogether. The racemose organ is well-marked though unlobulated or
only faintly so.

652 R. C. PUNNETT.

Collar. The musculature of the collar, like that of the proboscis, is somewhat feebly
developed. The cavity in the anterior part of the collar is very spacious. The intra-collar
lumen of the csophagus is unusually large (Pl. XLV. fig. 42). The dorsal collar septum
is present as usual behind the first root. The ventral septum may be found at the hind
end of the collar or it may be absent.

TABLE 4.

Pt. flava, var. gracilis.

The collar cord exhibits a continuous lumen throughout.

No. of specimen (1) (2) (3) (4) (5)
Proboscis coelom.......-...5+.c00s-0- Fairly large Not large as (2) Spacious as (1)
Collanicoclomisr-e-.s-4s-seereccescene Spacious as (1) as (1) as (1) as (1)
Ventral proboscis septum......... Nearly to tip as (1) as (1) as (1) as (1)

of stomochord
Dorso-ventral proboscis muscles Present as (1) as (1) — =
| Dorsal collar septum ............... Complete after as (1) as (1) as (1) as (1)
Ist root
Ventral collar septum ............ Traces only as (1) Absent as (3) Complete at
hind end of
collar
RACEMOSCIOLPAM! .0c.ccrcc.--eessesces Well marked | Well marked. | Well marked. as (1) as (1)
but small and | Unlobulated Slightly
unlobulated lobulated
Dorsal proboscis muscle decus- Fairly well Not very Well marked. | Very feeble. as (1)
sation marked. No | marked. No | No ventral cir-| No ventral cir-
ventral circu- | ventral circu- | cular muscles | cular muscles
lar muscles lar muscles :
Worsalll Toobs\scc. -scceccascesseuvs'esdees *RR RRr RRR RRr RRr 4
IPRODOSCISNPONES IN. ecessesessesenee ses *IR Lr IR IR L only
INGYVEXCOLG! weecctoescscteesssvaessnes Lumen con- as (1) as (1) as (1) as (1)
2 me ¥ oa tinuous
ength of post-brance ial groove ; , ; : ‘
- Length of branchial region - au oe ao i te
Length of cornua x 100 a } ; mee ‘
inept oneallan ns es 17-4 19:0 11:3 155 14:1
coe of branchial ces 1-26 114 1-00 60 85
Circumference of cesoph. ces.
Symapiiculaysssss-ssscsscesse-scacccses Not more than} Not more than | Not more than — Not more than
9 u 7 7

Three roots

are usually to

be found. The cornua of the nuchal skeleton are rather shorter than usual, extending over

about 1—1

of the collar length.

* See foot-note on p. 645.

THE ENTEROPNEUSTA. 653

Trunk. The branchial region in this variety is very short, averaging only a quarter
as much again as the collar length (see Table 12, p. 673). In the Maldive specimens
the branchial part of the cesophagus is at least as large and generally somewhat larger
than the ventral portion. In the Minikoi specimens on the other hand it is somewhat
smaller, a feature in which they approximate to laccadivensis. The post-branchial groove is
very short in the specimens from Hulule, not being more than about #, of the length of
the short branchial region. In the Minikoi specimens it is rather longer. The genital
pleurae are large in the branchial region (Pl. XLVI. figs. 50, 54). They diminish in size
considerably in the region of the post-branchial groove (Pl. XLVI. figs. 53, 55), after which
they are very small.

The proportions of the various regions of the body in the above account have been
determined from an examination of 28 specimens from Hulule (vide Table 12, p. 673).
Five specimens only were procured from Minikoi and, as will be seen in Table 4, Nos. 4
and 5, these differ somewhat from the Hulule forms. It is possible that examination of a
larger amount of material may shew that these Minikoi specimens belong to a distinct variety.
The difference in their mode of occurrence (p. 651) would seem to point the same way,
but in any case they are very closely related to the Hulule form.

Ptychodera flava, var. muscula (Pl. XLII. fig. 18).

Locauity, ETc. A Maldive form from Goifurfehendu Atoll, Goidu. It was obtained from
the sandy shore to the east of the island in the sand below the weed and under some
square slabs of beach sandstone. In company with it was found Spengelia porosa. About
a dozen specimens procured in all.

EXTERNAL FEATURES. Colour in life pale yellow with very black hepatic region. A
small form with a total length after preservation of 31—48 mm. Average length compared
with collar as unity = 156. A somewhat short and stoutly built form, generally speaking.
The anterior hepatic caeca are, after preservation, darker than the rest.

INTERNAL STRUCTURE.

Proboscis. The internal longitudinal musculature is very strongly developed and closely
surrounds the central complex, almost obliterating the proboscis cavity. No dorso-ventral
muscles anterior to the pericardium. The ventral proboscis septum is short and does not
reach nearly to the tip of the stomochord. Of the paired proboscis pores one may not
be functional. The racemose organ may be small or fairly well marked and slightly lobulated.
The nuchal skeleton has a well-developed keel which gives off a flattened expansion over
the racemose organ. Such an expansion occurs generally in the genus but is not usually
so well-marked as in the present variety (Pl. XLII. fig. 18). A strong dorsal muscular
decussation occurs in the anterior portion of the proboscis, and from it pass some circular
fibres ventrally.

Collar. The musculature of this region also isewell developed. The cavity in the
anterior part of the proboscis is comparatively large. The dorsal septum is found as usual
after the first root though it may not immediately be complete. The ventral septum is
absent or traces of it only are to be found.

The lumen of the nerve cord shews a marked tendency to become occluded. Three or
four dorsal roots may be present.

Gan. 84

654

Pt. flava, var. muscula.

R. C. PUNNETT.

TABLE

No. of specimen

(1)

IBrobOScis|COClOM sees. -caceeceseesete

Collar coelom

Dorso-ventral proboscis muscles

Dorsal collar septum

sede eeeweees

Ventral collar septum

Racemose organ

Dorsal proboscis muscle decus-
sation

Dorsal roots

Proboscis pores

Nerve cord

Length of post-branchial groove
Length of branchial region

Length of cornua x 100
Length of collar

Circumference of branchial ces.
Circumference of cesoph. ces.

Synapticula

Length of branchial region
Length of collar

Very small
Spacious
Not nearly
to tip of
stomochord
Absent
After 1st

root. Not
complete

Traces

Very small

Strong with
ventral cir-
cular fibres

*RRR
*Lr
Lumen with

tendency to
obliteration

14:0

| (2)

| Small

Fairly large
as (1)

as (1)

After 1st
root complete

Absent

Well marked.
Slightly
lobulated

as (1)

RRrr
LR

as (1)

14°5

15°8

1:30

5.

Pt. flava, var. cooperi.

(1)

Spacious

Very
spacious

Not nearly
to tip of
stomochord
Absent

Complete
after 1st root

Absent

Well marked.
Unlobulated

Feeble. No
ventral cir-
cular muscles

IR
Lumen with

tendency to
obliteration

“12

119

Not more
than 8

2°23

Pt. viridis.

(2) (1) (2)
Small Very small Small
as (1) Small Very small
as (1) | Not nearly | Not quite

to tip of to tip of
stomochord | stomochord
as (1) Absent as (1)
as (1) Complete After Ist
after Ist root| root but not
at once
complete
Traces in || Complete at Traces
extreme hind|/extreme hind
end of collar || end of collar
as (1) Large. Large and
Unlobulated | _ slightly
lobulated
Moderately 2 Strong with
developed. a few ventral
No ventral fibres
circular
muscles

Rr RRr 2

R 2 LR
as (1) Lumen as (1)

completely
obliterated
Very short
‘15 and insigni- as (1)
ficant
53°7 161 27°8
1:13 1:02 95
Not more = =
than 7
2°06 “85 1:44

* See foot-note on p. 645.

THE ENTEROPNEUSTA. 655

The cornua of the nuchal skeleton are short, extending over less than } of the collar
length.

Trunk. The branchial region is fairly long, averaging more than double the length of
the collar in 10 specimens (see Table 12, p. 673). In one small specimen only was it
less than double the length of the collar. On the whole it is remarkably constant in length.
The branchial division of the cesophagus is considerably less than the ventral portion. The
post-branchial canal was found to be short in the specimen examined, being less than } of
the length of the branchial region. The post-branchial pre-hepatic genital tract is compara-
tively short. The genital pleurae are well-developed.

Ptychodera flava, var. cooperi (Pl. XX XVIII. fig. 12; Pl. XLV. figs. 45—4/7).

Locatity, ETc. Two specimens, both somewhat imperfect, from N. Male Atoll. Dredged
in 5 fathoms on a sandy bottom within the velu of Jaro near Helengeli.

EXTERNAL FEATURES. A small form with a maximum breadth in the collar and branchial
region of less than 3mm. Behind the hepatic region the diameter of the animal is only
a trifle over 1mm. In the preserved specimens the collar and proboscis are both very short,
their width being greater than their length (Pl. XXXVIII. fig. 12). The genital pleurae
are large in the branchial region, which is rather more than double the length of the collar.
The genital region between the termination of the branchial and the commencement of the
hepatic caeca is relatively long. The hepatic caeca are short but well-developed, and on
their inner sides are lobulated. There is no distinction into lighter and darker ones.

INTERNAL STRUCTURE.

Proboscis. The cavity of the proboscis is well-developed in one specimen, whilst in
the other it is small. The radial arrangement of the muscles is not so conspicuous as in
most members of the genus. Dorso-ventral muscles anterior to the pericardium are not
present, and the ventral proboscis septum is very short. The right proboscis pore in each
case places the dorsal coelom in communication with the exterior. The left pore is present
in one specimen but absent in the other. The racemose organ is well-marked but unlobulated.
The keel of the nuchal skeleton is somewhat larger than usually obtains in the species
(Pl. XLV. fig. 45). The skeleton is altogether massive (Pl. XLV. fig. 46).

Collar. The cavity in the anterior part of the collar is very spacious. Posterior to
it the collar musculature is unusually strongly developed, a fact which may in some measure
account for the extreme shortening of this region on preservation. The dorsal septum is
as usual complete after the first root. A ventral septum was completely absent in one
specimen though traces of it occurred at the extreme hind end of the collar in the other.
In one case a single dorsal root reaching to the epidermis was found. In the second
specimen this was supplemented by a rudimentary one. The lumen of the collar cord shews
a tendency to occlusion behind the level where the roots come off.

A noteworthy feature of this variety is the condition of the nuchal cornua which extend
back to the hind end of the collar, the tip of them being found in sections which also
pass through the first branchial bars. Such a condition is most unusual amongst the
Ptychoderidae, recalling that recently described by Ritter (OO, p. 113) for Harrimania
maculosa.

Trunk. The branchial portion of the cesophagus is rather larger than the ventral portion.
Seven to eight synapticula are present. The post-branchial groove is large (Pl. XLV. fig. 47),
84—2

656 R. C. PUNNETT.

whilst the cesophagus at this level is very small. The genital pleurae are still large in this
region but become much reduced shortly after it.

Ptychodera viridis, n. sp. (Pl. XXXVII. figs. 2, 6, 7; Pl. XXXIX. fig. 32; Pl. XLII.
figs. 17 and 19).

Locatity, ETc. From Hulule, Maldive Is. Three specimens only, two being complete,
were dug from sand towards the cast of the island.

EXTERNAL FEATURES. Coloured sketches from life of this worm were fortunately made
by Mr Forster Cooper. From the reproductions of these on Plate XXXVII. it will be seen
that green is the prevailing tint on the body of the animal, though the proboscis is pale
yellow, and the collar pale yellow with a certain amount of orange. To the predominant
body tint the animal owes its specific name. From the sketches it appears that the proboscis
is considerably longer than the collar, a proportion which still exists in the preserved creature.
The length is not great. Of the two complete specimens one measured 36 mm. and the
second only 21 mm. in length after preservation (ratio of total length to collar length being
respectively 18:1 and 14:1). Compared therefore with such forms as Pé. flava, var. lacca-
divensis, this species is a small, short, and somewhat stunted form. The genital pleurae
are well-developed. The external liver saccules are relatively feebly developed and are of
a uniform colour.

INTERNAL STRUCTURE. The musculature of the proboscis is strong and compact, the
radial bundles into which the longitudinal muscles are gathered being closely connected by
connective tissue. The powerful development of the longitudinal muscles leads to the practical
obliteration of the proboscis cavity (Pl. XXXIX. fig. 32). A well-marked dorsal muscular
decussation occurs in the anterior part of the proboscis (Pl. XX XIX. fig. 32, dd) and
fibres from it pass ventral to the central complex. Dorso-ventral muscles anterior to the
pericardium are not present. The ventral proboscis septum reaches forwards almost to the
tip of the stomochord. The racemose organ is small and unlobulated. Both proboscis pores
are present though only one is in functional communication with the dorsal proboscis coelom.
This may be either the right or the left one.

Collar. The musculature and the connective tissue are here strongly developed and
the cavity of the collar is very much reduced. The dorsal septum occurs after the first
root as usual and is generally complete. Traces of the ventral septum may occur. The
lumen of the collar cord is almost entirely obliterated throughout and the cord in section
has as great a dorso-ventral diameter as a lateral one (PI. XLII. fig. 19). In one of the
specimens sectioned three roots were present of which the last was rudimentary.

The cornua of the nuchal skeleton apparently vary much in length. In one specimen
they extended over about } of the collar, whilst in another they were rather more than a
quarter as long as this structure.

Trunk. The branchial region is short. Compared to the collar length as unity it
measured in the three available specimens ‘85, 1:00, and 1°44, giving an average of 1:10.
The branchial portion of the cesophagus is approximately of the same size as the ventral
portion. The post-branchial groove is very short and insignificant. The genital folds are
large in the branchial region and also in the region of the post-branchial groove (Pl. XLIL
fig. 17).

THE ENTEROPNEUSTA. 657

Ptychodera asymmetrica, n. sp. (Pl. XX XVII. figs. 1 and 9; Pl. XLVI. figs. 52, 56, 58).

Locauity, Etc. From Hulule. According to Mr Stanley Gardiner, “Near the south islet
is a pool with loose rocks well within the boulder zone. This form was found here in
accumulations of sand under stones. They are remarkable in life for the enormous quantity
of mucus they secrete, so much indeed that it is almost impossible to obtain them clean
for preservation.” About 10 specimens were procured, mostly somewhat fragmentary.

EXTERNAL FEATURES. A small form measuring on the average about 40 mm. in length
(Pl. XXXVIT. fig. 1). One larger specimen, a sketch of which was made by Mr Forster
Cooper and is reproduced on Pl. XXXVII. fig. 9, measured after preservation 60 mm. in
length with a collar 3mm. long. This specimen however was unusually large. Two points
about its external appearance merit attention. In the first place the liver saccules are
somewhat small and uniformly dark in colour. There is no sharp line of demarcation between
an anterior set of very dark caeca and a posterior set of lighter ones as in Pt. flava, var.
laccadivensis. In the second place careful examination shews that the left genital pleura is
always somewhat larger and more swollen than the right one. In all the eight specimens
examined by me the gonads shew asymmetry by being developed only on the left side.
This was the case both in a very young specimen with quite immature gonads, and also
in the large specimen above alluded to im which the gonads were full of ripe spermatozoa.
In his account of Pt. flava Willey (99, p. 240) mentions a case in which the gonads
were only developed in the right genital pleura, those in the left being apparently in a
state of arrested development. Evidently with the case of Asymmetron in his mind Willey
writes, “Such a differential behaviour of the two sides of the body is of interest as indicating
a tendency to unilaterality in the matter of the gonads.” Spengel (O83, p. 305) criticises
this remark of Willey’s, regarding such a condition as produced by the presence of a
parasitic copepod, Zve sp. In the two cases examined by Spengel where the gonads of one
side were undeveloped the parasite occurred in the genital pleura of that side, and there
seems little doubt that Spengel’s explanation is here correct, and that the condition is a
pathological one. But there can be no doubt that this explanation will not hold for Pt.
asymmetrica. The fact that in eight cases, in different stages of growth, gonads were always
absent from the right pleura, coupled with the fact that no parasite was to be found,
seems to shew beyond all question that unilaterality of the gonads is a feature which is
characteristic of this species.

INTERNAL STRUCTURE.

Proboscis. The longitudinal muscles do not exhibit a markedly radial arrangement of
bundles. They are strongly developed and almost entirely fill the cavity of the proboscis
so that there is no space, or only a very small one, between them and the central complex.
There are no-dorso-ventral muscles in front of the pericardium. The ventral proboscis septum
does not reach to the tip of the stomochord and is often much shorter.

The racemose organ is subject to considerable variation. It may be small and unlobulated,
large and lobulated or not.

There is usually a well-marked dorsal muscular decussation in the anterior part of the
proboscis though no circular fibres pass from it ventrally.

Collar. The cavity of the collar varies considerably. It is on the whole not large
and may be almost absent. The collar musculature is well-developed. The dorsal septum

658

R. C. PUNNETT.

is always complete after the first root.

The ventral septum is a more conspicuous structure
in this species than in most others of the genus. It is always found in the posterior part
of the collar and may reach forwards over 4 of the collar length.

TABLE 6.

Pt. asymmetrica.

No. of specimen (1) (2) (3) (4) (5)
Proboscis coelom............+-s-+e+-« Very small as (1) Well marked as (1) Obliterated by
but small muscles
Collanicoelom\cennceas-sesrssesseoeee Much reduced Almost Fairly spacious} Very small Well marked
obliterated
Ventral proboscis septum ......... Short. Not | Not to tip of | Not quite to as (3) as (1)
nearly to tip of | stomochord tip of
stomochord stomochord
Dorso-ventral proboscis muscles Absent as (1) as (1) as (1) as (1)
Dorsal collar septum ............... Complete after as (1) as (1) as (1) as (1)
Ist root
Ventral collar septum ............ Complete in as (1) as (1) as (1) Complete in
hind end of posterior 4 of
collar collar
RaCemose Oran) ..-..<.--tasssceeeres Large but Large and as (1) Small and | Very large and
unlobulated lobulated unlobulated lobulated
Dorsal proboscis muscle decuss- | Well marked. as (1) as (1) Not well as (1)
ation No ventral cir- marked. No
cular muscles ventral circular
muscles
Dorsal \OObS ss sneseresse secre ome eceeee *RRR RRrr RR R RR
IBrOHOSCIS POLES eecpess-cseessree eee *LR Lr 1R IR IR
IN Gives Cord: hr scerss- ses cocs-ose meets Lumen almost |Lumen distinct| Lumen distinct as (1) as (1)
obliterated in places in places.
Strong tendency
" to obliteration
ength of post-branchial groove Bs , .
Length of branchial region ou ay ee um
Length of cornua x 100 : ! :
Reaeiioncallaae ae 245 20°9 47°3 40°5 25°7
Circumference of branchial ces. 98 1-08 1-04 -s 1-06
Circumference of cesoph. ces.
SYMAP HCW cesecssstsecsosesceseses -- -- Not more than 5 _— =

The lumen of the collar cord is never continuous throughout. It may be present as
a number of small cavities much broken up, or it may be practically absent, the cord shewing

* See foot-note on p. 645.

THE ENTEROPNEUSTA. 659

only the merest traces of a lumen. As is often the case with the collar cord, when the
lumen shews a marked tendency to occlusion, the dorso-ventral diameter as seen in transverse
section tends to become relatively great as compared with the transverse diameter (Pl. XLIV.
fig. 29). The number of the dorsal roots is very variable. There may be as few as one,
though four seems to be the maximum.

The cornua of the nuchal skeleton are on the whole long for members of this genus,
but shew a considerable amount of variation. They may extend over only 4 of the collar
length, or their length may be equal nearly to } that of the collar.

Trunk. The branchial portion of the cesophagus is of about the same size as_ the
ventral part. The relative length of the branchial region as compared with the collar length
is short, it being seldom more than half as long again as the latter and sometimes even
less. In the large specimen figured on Pl. XXXVIL. fig. 9, however, it was rather more
than three times the collar length.

The post-branchial groove on the other hand is longer than usual, averaging between
and 4 the length of the branchial region.

on

The genital folds are somewhat small (Pl. XLVI. figs. 52 and 56) and become mere
ridges in the region of the post-branchial groove (Pl. XLVI. fig. 58).

SOME ANATOMICAL POINTS.

As Spengel in his latest pubheation (08) differs on one or two points from Willey
it may be worth devoting a few lines to them. In two of these cases, the pygochord and
the lateral septa, I find myself in agreement with Spengel rather than Willey.

The pygochord is a structure on which Willey has laid some stress, describing it as
“a longitudinal, solid, supporting band” (99, p. 243), and evidently seeing in it a skeletal
structure for the support of the caudal region. On the ground of its minute proportions
Spengel (O08, p. 317) dissents from Willey’s interpretation, and one cannot help agreeing
with Spengel in refusing to regard such a small and rudimentary structure as of any
importance from this point of view. As Spengel remarks we have absolutely no knowledge
of the function of this apparently vestigial structure, and it is with some diffidence that
I venture to suggest that it may be the remains of a ventral siphon which was at one
time of functional importance but now vestigial. The occurrence of such a siphon, whatever
may be its exact function, is common among sand-feeding animals. Similar structures occur in
the Echinoidea, in the Capitellidae, and in certain Chaetopoda such as Thallasema and Echiurus
—all creatures which derive their nutriment from among the large quantities of sand con-
tinually passing through the alimentary canal. The condition of the pygochord found in a
specimen of Pt. saxicola is interesting in this connection. The structure in question was
here connected at either end with the intestine whilst for the middle part of its course it
was entirely free (cf. Pl. XLIV. figs. 835—37), a condition strongly suggestive of a collateral
intestine.

The lateral septum is described by Willey as passing to the epidermis at the base
of the genital pleurae in the post-branchial region (99, Pl. XXIX. fig. 14). Spengel,
however, finds that this is not so, but that it ends at the lateral blood vessel (O38, p. 297).
It is sufficient to remark that on this point I find myself in agreement with Spengel.

660 R. C. PUNNETT.

Lastly there is a point in connection with the branchiae which calls for short notice.
It has been assumed that the main function subserved by these structures is that of
respiration, and in the figures given by most authors the blood vessels form a prominent
feature. Nevertheless the course of the branchial circulation is a point which has never
been satisfactorily cleared up (cf. Delage and Herouard, ’98, p. 41), and in looking through
numerous series of sections through the branchial region I have been struck by the infrequency
with which one finds any blood in these structures. On the other hand the sub-epidermal
vascular plexus is exceedingly well-developed everywhere and it is difficult to avoid the
conclusion that respiration is mainly, if not entirely, carried out by the large skin area, whilst
the main function of the gills is to act as a sieve for straiming off the water from the
mixture of sand and water continually swallowed by the animal.

THE FORMATION OF THE GONADS.

The origin of the gonads is a subject on which widely diverse views have been upheld
by different observers. Bateson (?86) attributed to them an ectodermal origin, Spengel (795)
regards them as being developed in a blood space, whilst Morgan (?94, p. 60) holds that they
are formed from the mesodermal elements of the coelom. The question is one of considerable
difficulty and none of the above observers has made out a very strong case in support of
his view. Bateson bases his belief on the early connection apparent between the gonad and
the ectoderm (86, Pl. XXXII. fig. 110). Spengel considers this connection to be due to
the precocious formation of a duct and claims to have found sex cells at a very early
stage of development inside the blood vessels (95, Pl. II. figs. 27 and 28). He finds
additional support for his view in the fact that in a small specimen’ of Pt. flava, var.
laysanica, the “Anlage” of the gonad first appears between the two lamellae of the lateral
septum (’08, p. 302). The case which he has made out is far from convincing since it
is by no means certain that the small cells which he regards as primitive sexual cells are
of that nature, whilst on the other hand there are good grounds for looking upon the
lateral septum as composed of four rather than of two lamellae (see below, p. 661). Lastly
Morgan’s contention as to the mesodermal origin of the gonads is not borne out by his
figures. In the earliest stage figured by him (94, Pl. VI. fig. 79) the gonad is obviously
im continuity with the ectoderm, a connection which, to judge by his figures, is very soon
lost, though doubtless re-established later on the formation of the genital ducts. The question
of the ectodermal origin of the gonads turns largely upon the interpretation of the ecto-
dermal connection found at different stages of their development. In other words, has the
gonad an ectodermal connection at two different stages of its growth, the earlier representing
the ingrowth of ectoderm to form the gonad, whilst the latter represents the functional
duct? From the following observations on Pt. flava I am inclined to believe that such is
the case. As the gonads appear to develop earlier in some varieties than in others I have
considered them apart with the following results.

(1) Var. laccadivensis. The youngest stages of the gonads were met with in a very
small specimen about 15 mm. long. ‘The young gonads are in many places in connection
with the ectoderm (Pl. XL. fig. 36). In the region of the genital pleurae this connection

1 Judging from the regenerated material at my disposal in doubt in considering this small specimen of Spengel’s to be a
the case of Pt. flava, var. laccadivensis, I have very little regenerated one.

THE ENTEROPNEUSTA. 661

is almost invariably found where the lateral septum joins the basement-membrane at the
tip of the pleura. Occasionally an ingrowth from the ectoderm of the inner surface of the
pleura is to be seen, but this is rare. The irregular occurrence of the ectodermal connections
and their somewhat attenuated condition seems to indicate that they are in process of
disappearance. And this supposition is borne out by the fact that in somewhat older
specimens the gonads are devoid of any ectodermal connection.

(2) Var. sawicola. Though all the specimens of this variety were of fair size the gonads
in all are in a very backward state of development. The gonads are in all cases young
and in all cases possess a more or less well-marked ectodermal connection. This connection
is relatively wider in the younger gonads (cf. Pl. XL. figs. 41 and 42). A still younger
stage of an ectodermal ingrowth at the tip of a genital pleura is shewn on Pl. XXXIX.
fig. 31. There can be little question but that this is of the nature of an ingrowth, and
indeed rounded cells may be seen in it, which are not to be found in other parts of the
ectoderm, and which are probably to be regarded as primitive genital cells. The main
ingrowth is that at the tip of the pleura as in var. laccadivensis, but the accessory ingrowths
on the inner surface of the pleurae are much more numerous in saaicola, a fact probably
correlated with the larger size attained by the animal for a similar stage of gonidial develop-
ment. Unfortunately none of the four specimens of this variety possessed older gonads.

(3) Var. muscula. In a specimen with immature and somewhat small gonads an-ecto-
dermal connection is generally wanting. It may be present here and there. In an older
specimen in which spermatocytes could be recognized in the gonads (Pl. XL. fig. 37) the
ectodermal connections are present on almost all the gonads.

(4) Var. maldivensis. This variety is characterized by often having long slender genital
ducts. They are found connected with fairly well-developed gonads and in some cases I
have been able to make out their double origin, in part from an ectodermal ingrowth and
in part from an outgrowth from the gonad. In one or two instances the two portions had
not effected a junction, a condition supporting the view that the true genital ducts are
late in making their appearance. It has already been noticed that the ectodermal ingrowths
near the tip of the pleurae always occur directly over the point where the lateral septum
joins the basement-membrane and pass down between its lamellae. On the other hand the
ducts in connection with the gonads nearest the tip of the pleurae may be entirely inde-
pendent of the lateral septum (Pl. XX XIX. fig. 27).

All the above observations are in harmony with the view that the gonads are (in
Pt. flava at any rate) derived from ectodermal ingrowths, that they subsequently lose their
connection with the ectoderm, but regain it later when the genital ducts are established.
Further, on this view we can explain the lateral septum of the Ptychoderidae, regarding it
as having in the first place been brought about by the basement-membrane being carried
in by the primitive ectodermal ingrowths at the tip of the pleurae (at that time probably
very much smaller). On this view the lateral septum should be four-layered since the
basement-membrane consists already of two layers enclosing blood spaces. That four potential
layers are present is shewn by the fact that the apparently single layer round the gonad
is in reality a double one since blood spaces frequently occur in it (Pl. XL. fig. 41). The
cavity of the gonad is therefore quite distinct from the blood EES, which militates against
Spengel’s view of the nature of the gonad.

The gonads of Ptychodera flava are characterized by the presence of peculiar bodies
G. IL. 85

662 R. C. PUNNETT.

apparently devoid both of structure and of nucleus. The origin of these bodies, the “ fettartig
Kiigelchen” of Spengel and the “eosinophil globules” of Willey, is somewhat obscure. Willey
holds that they are the products of nuclear degeneration accompanied by hypertrophy of the
nucleolus. Spengel (?O8, p. 306) refuses to believe that they are the outcome of degenera-
tive changes. In the young specimen of Pt. laccadivensis referred to above I have been
able to follow what seems to be the history of these bodies. Certain of the spindle-shaped
mesodermal cells become first oval and then more or less spherical in shape (PI. XL.
fig. 38, a—b), at the same time increasing slightly in bulk. A few granules make their
appearance in the hitherto clear cytoplasm. A little later these cells appear to collect
together in clusters of about 6—10 (fig. 38, c), which subsequently fuse together to form
giant cells packed with granules and with the nuclei degenerating (fig. 38, d and e). Later
the nuclei disappear and the giant cells apply themselves close to the gonad. Where this
occurs the wall of the gonad seems to break down and by some method which is not clear
the nutriment stored in the giant cells is transferred to the interior of the gonad where
it loses its granular appearance (cf. Pl. XL. fig. 37). At later stages small round corpuscles
of an eosinophil nature also make their appearance in the mesoderm and subsequently apply
themselves to the gonads in a similar manner (Pl. XL. fig. 41). The chief interest of the
process seems to lie in the fact that although the sexual cells themselves are derived from
the ectoderm’, it is the mesoderm which provides them with nourishment and which con-
tributes the yolk to the eggs.

The above account is of interest when it is considered with reference to our conception
of the nature and properties of the coelom. According to present ideas one of the chief
functions of the coelom is to give rise to the generative cells from the epithelium which
lines it. Further it is usually held that the most primitive method of formation of the
coelom is that of archenteric diverticula. In animals like the Enteropneusta, where this
method of the formation is found, we should naturally look for genital cells arising from
its lining. That this is not so may cause us to regard with some reservation the statement
that the coelom was originally a gonocoel’, and to consider it rather as an organ specialized,
among other things, for the reception and maturation of the genital cells.

VARIATION IN PTYCHODERA FLAVA.

The different varieties of Pt. flava are by no means easy to distinguish from one another
after preservation. An adequate examination of the characters which are of systematic value
involves the preparation of a large number of serial sections. If a trustworthy criterion based
upon external features alone could be found much labour might be spared the Enteropneustic
systematist. The large amount of material collected by Mr Stanley Gardiner has enabled
me to attempt this.

The characters upon which such a criterion might be based must satisfy two conditions;
(1) they must be capable of easy measurement, and (2) they must be independent of the

1 In this connection it is interesting to notice thatsolong figs. 88 and 89).
ago as 1885 Hubrecht described the ectodermal origin of the 2 For the most recent general account of the coelom the
gonads in the Nemertean Lineus gesserensis. He was able reader may be referred to Lankester’s Treatise on Zoology,
here to distinguish with certainty the primary ectodermal 1900, Pt. 1. chap. 11.
connection from the later formed duct (Hubrecht, ’85, PI. vy.

THE ENTEROPNEUSTA. 663

growth of the animal. This latter condition precludes the direct use of absolute measurements.
It is obvious that a ratio must be made use of—a simple ratio whose two factors are
equally affected by growth changes. Though several distinct regions may be distinguished
in the body of a Ptychodera, with the exception of the collar and branchial region, their
limits are not very clearly defined. Accordingly most attention has been paid to the ratio
of branchial to collar length. And this more especially since any collection of Enteropneusts
will be largely made up of damaged specimens, and though few specimens may attain to
anything like completeness, yet in the great majority both collar and branchial regions will
be perfect. Great accuracy in the measurement of such animals as the Enteropneusta is
out of the question since in a soft-bodied creature relative differences in’ the state of
contraction of the various body regions may easily occur, whilst the personal equation must
enter largely into the result. In the present instance however all the material had been
treated in the same way, ze. had been narcotised in chloral hydrate for several hours before
preservation, thus ensuring an equable contraction, whilst the personal equation was as far
as possible eliminated by my making all the measurements within the space of a week or so.

By far the most frequent of the varieties described in this paper is laccadivensis, and
it is here that the measurements are most complete. In addition to the length of the
collar and of the branchial region the total length and the length of the genito-hepatic
regions was measured wherever the specimen was sufficiently perfect. The width of the
collar was also registered. The results for 123 specimens (fewer in the case of the genito-
hepatic and total lengths) are given in Table 11, pp. 671—2, and in the diagram on p. 666.
Before however any value can be attached to these figures it must be shewn how, if at
all, growth affects these ratios.

The method employed for estimating growth changes was to divide the animals into
several groups according to size, and to calculate the above three ratios for each group
separately. As the collar width was sometimes less, sometimes greater, than the length it
was thought advisable to take this factor also into account, and in the case of laccadivensis
the following groups were made.

Numbers in Group
Branchial Genito-hepatic Total

Group 1. Collar length x breadth not >16 ............... 47 38 19
m 2b aS ty) PRS > 16, but not > 25... 45 33 17

5 Ok ”» » ” SP, sy py Se blshes 27 15 13
sree * fie oe of Sev eciea atic aa eeicichis ebesinie= 4 —- _-
123 86 49

The mean of each ratio was then caleulated for each of the above groups with the

following result :—

No. of Specimens
Branchial length

settee ee Ne 0 = 2°08 47
Colle Iocan Mean for Group 1 0: 7
” ” ” 2 = 218 45

” ” ” 3 = 2°36 27

4=277 4

Average 2°20

664 R. C. PUNNETT.

No. of Specimens
Genito-hepatic length

é nek 4 (Gk Parole :
Collar length Mean for Group 1 = 4:94 38
” ” ” 2 = 4:90 33

” » ” 3 = 484 15

Average 4:90
Total length

Gollanilerent? Mean for Group 1 = 23:1 19
” ” ” 2 = 22:4 ile
” ” ” 3 = 23-4 13

Average 22°9

An examination of the above table brings out clearly the important fact that two of
the three ratios, ie. those of the genito-hepatic and total lengths, are quite unaffected by
growth changes, whilst that of the branchial region is not greatly affected. Neglecting
group 4 in which there are so few specimens there is a gradual rise in the ratio with
increase in size from 2°08 in group 1 to 2°18 in group 2, and 236 in group 3. There is
therefore an increase of about 12°/, in group 3 as compared with group 1, an increase
sufficiently well-marked though not very great when we take into account the unavoidable
roughness of the measurements. A possible source of error must be noticed here. A con-
siderable number of specimens (some 8—10°/,) were regenerating. Where this process was
obviously at work the specimens were not mcluded in the above measurements. In the
latest stages, however, regenerated individuals are impossible to distinguish from ordinary
ones with a somewhat shorter branchial region than usual. As regeneration is far more
common in the smaller specimens it seems almost certain that the values of the ratio for
groups 1 and 2 are somewhat too low in comparison with those of group 3. On the whole
it would appear from these measurements on laccadivensis that the proportions of the body
of this Enteropneust are very little, if at all, Hable to alteration during the later stages
of growth.

This conclusion is borne out by a similar series of measurements made on Pt. flava,
var. maldivensis. The 52 specimens of this variety were divided into three groups of which
the last contained only the five largest specimens (see Table 13, p. 674). On taking the
mean for the three ratios in each group the following figures were obtained.

TABLE 7.
No. of Specimens
Branchial length
ey Mesnetorran = 165
Collarslencth Mean for Group 1 = 1 19
” ” ” 2 = 1:62 28
Ao nS » & = 2:00 5

Average 1°67
Genito-hepatic length

Gallarmeneth Mean for Group 1 = 4°58 19
” ” ” 2= 414 28
” ” ” 3 = 5:23 5

Average 4°41
Total length
Collar length’

Mean for Group 1 = 165

” ” ” 18°7
” ” ” 3 = 17°6 2

Average 17°7

bo
ll

THE ENTEROPNEUSTA. 665

There is more irregularity here in the figures for the genito-hepatic length and_ total
length, which perhaps is only to be expected in view of the smaller number of specimens.
There is however no uniformity indicating a growth change. The value for group 3 in the
branchial series is far larger than for groups 1 and 2. This however is due to the large
branchial region of a single specimen (Table 13, No. 49) and its effect on the few individuals
of group 3. The closeness of the figures for groups 1 and 2 (1°65 to 1:62), where there
are more specimens available, points to the ratio of branchial length to collar not being
affected by growth changes. Assuming then, and the assumption seems a fair one, that
growth has little or no effect on the relative proportions of the body in Ptychodera we have
before us an easily applied criterion for distinguishing different positions of organic stability
in this genus. We have but to determine one of these ratios (and in practice the branchial
one will be found the most convenient) on a number of specimens of each supposed variety,
and to apply to the series so obtained the ordinary biometric methods for measuring the
variability. Marked differences occurring in two groups of specimens would at once cast a
doubt on the identity of two such groups.

This method has been adopted here for such of the varieties as possessed nine or more
specimens. The branchial ratio alone was used owing to the fragmentary condition of most
of the individuals in each variety. On working out the mean and standard deviation in
each case the following results were obtained :—

TABLE 8.’
Pt. laccadivensis | Pt. maldivensis | Pt. muscula Pt. gracilis Pt. parva |
| (Minikoi) C4 (Goidu) (Hulule) (Turadu)
a —— — —| es = = —
M. 2263 1-630 | 2°225 _ 1259 1514
P. E. M. + 1405 + 1724 } + “2484 + +1995 1753
o + 2°3098 | +1:8431 | +1:1647 . +1°5656 7798
P. E.o + 0993 | + 1219 iO es Neti “1240
C.V. 102°07 | 113°08 52°35 124°35 51°50
N. 123 52 | 10 28 | 9

These results shew clearly that in certain cases we are without doubt dealing with different
races’ of Pt. flava. Thus the two forms laccudivensis and maldivensis are exceedingly alike
both in external appearance and in internal structure. The branchial ratio is however
very different, the mean for maldivensis (1°63) being very much lower than that (2°26) for
laccadivensis. And the difference between the two is almost four times the probable error of
either, from which it may be calculated that the odds are many thousands to one against
its being due to errors of random sampling alone.

! For the actual measurements on which these figures are
based see Tables 11—13, pp. 671—4. Of the letters in the

(ef. Davenport, C., Statistical Methods, 1899, p. 38). My
point is that in the group of animals collectively designated

above Table (i.e. Table 8), M.=mean; P.E.M.=probable
error of the mean; o=standard deviation; P.E.c.=prob-
able error of standard deviation; C.V.=coefficient of varia-
tion; whilst N.=number of specimens in each case.

*I am not here concerned with the question of the
statistical method as a criterion of species and variety

Pt. flava there are to be found different positions of organic
stability, and in giving to these ‘‘a local habitation and a
name”’ there is no thought of distinguishing by statistical
methods between the terms, race, variety, and species. For
the value attached to such distinctions must for the present
be largely a matter of individual taste.

PUNNETT.

C.

R.

666

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THE ENTEROPNEUSTA. 667

In fact from these figures one may affirm with some confidence that laccadivensis and
muscula (though they cannot be separated from one another by this criterion) are quite
separate from maldivensis, gracilis, and parva, whilst of the last three gracilis is very
probably distinct from the other two. By this criterion alone it is not possible to separate
maldivensis from parva any more than laccadivensis from muscula. No doubt this could be
done in the case of these forms if the material were sufficient to treat another ratio (e.g.
the length ratio) in the same way.

The total length ratio was worked out for laccadivensis and maldivensis, in which forms
alone the amount of material rendered the attempt feasible. As there are no definite segments
by which to reckon, the absolute value obtaimed for o will depend upon the number of groups
into which the material is broken up. In this case the mean in the two varieties is 23'1 and
17-9 respectively. The series have been broken up into groups separated by an interval of
two units, which is approximately 10°/, of the mean in each case. The treatment has been
uniform in each case and the results are as follows :—

TABLE 9.

var. laccadivensis var. maldivensis

M. = 23-1 17-9
PE.M.=| + 129 + -260

c = +1°798 | +1°589 |
P.E.c= + -091 | + -184 |

Nee 49 | 17 |

Hence the mean is widely separate in the two forms—separated by so many times
the probable error as to render it absolutely certain that no random selection of 17 specimens
from a large quantity of lJaccadivensis could ever have a mean length ratio with the remotest
approximation to the above 17 specimens of maldivensis.

Leaving now the question of variation as a criterion of species and variety we may
pass on to consider one other point in connection with it. A great feature in the group
of the segmented Chordata is the process of cephalization which becomes more and more
pronounced as we pass upwards from Amphiozus to Mammals. Accompanying this process,
and doubtless intimately bound up with it, is the phenomenon of unequal variability in the
different regions of the animal’s body from the meristic pomt of view. With increased
cephalization we find associated a decreased variability at the cephalic as compared with the
caudal end. The nearer we approach to the caudal extremity the greater appears to become
the meristic variability’. From this point of view the relative variability of the different regions
of the Enteropneust’s body becomes of interest, especially as most writers see in the collar
cord the commencement of the cephalization process. From the data given in Table 11, p. 671,

1 For the present this statement is based upon a number dorso-lumbar and sacral regions a small amount, whilst the
of unpublished data from different Vertebrata, chiefly fishes, caudal region exhibits most of all. Probably some such
collected by the writer. The phenomenon is perhaps most phenomenon is to be found also among Invertebrates, but
familiar in the case of the Mammals where, speaking gener- absence of data permits only of conjecture.
ally, the cervical region exhibits no meristic variation, the

668 R. C. PUNNETT.

I have therefore estimated the variability of three separate regions of the body in Pt.
laccadivensis, viz. the branchial region, the genito-hepatic region, reaching from the hind end
of the branchial region to the point of junction of light and dark hepatic caeca, and the
hepatic-caudal, extending from this point of junction to the anus. In the following table
this is expressed quantitatively when the unit of variation’ selected was the same in all
three cases, 7.e. collar length x 1.

TABLE 10.
| | ayy ox 100
| | c | CV.=- Ste
|
| Branchial TESION' |. 7. .csse + ‘6712 | 29°66
Genito-hepatic region il +1°015 | 20°72
Hepatic-caudal region ...| +2°711 11-79

From this table it appears that the variability as measured by the standard deviation
gradually rises as we recede from the collar. But these three regions are of very different
length and the unit of measurement (collar length) bears towards the branchial region a
relation as regards length very different to that which it bears towards the hepatic-caudal
region. To avoid this difficulty we must consider the coefficient of variation (C.V.) in each
case, and when this is worked out (Table 10) it is at once evident that the relative variability
of the branchial region as regards length is far greater than that of the genito-hepatic,
and this again than that of the hepatic-caudal. In a word, as we proceed caudalwards from
the collar the relative variability of the body as regards length becomes less. In Amphiowus
and the Vertebrata on the other hand the relative variability of the body as measured by
the number of segments becomes greater as we proceed caudalwards. One of three alter-
natives would seem to follow:—either (1) we must regard the processes of differential growth
as widely different in the two cases, or (2) we must look upon the number of a meristic
series as altogether independent of its length, or (8) we must consider that there exists in
Ptychodera a somewhat vague segmentation—that the animal is composed of morphologically
equivalent though ill-defined segments of which the length differs considerably in different
regions of the body. The amount of favour with which we can regard the first of these
suggestions depends upon our conception of the relations of the Enteropneusts to the Chordata.
Those who are inclined to uphold the chordate affinities of this group would naturally turn
to either of the other two alternatives. The former of these I regard as untenable, since
such data as I have been able to collect on the subject all tend to shew that there is
a fairly high correlation between the relative total length of a vertebrate axial meristic
series and the number of units of which it is composed. There remains to be considered
the last of these hypotheses. Are there grounds for attributing any form of segmentation®
to the Enteropneustic body, and further, if this is the case, for considering that such
segments may differ in size in the different regions of the animal? To the first part of
the question the answer must be in the affirmative. In the branchiae, in the hepatic caeca,
and in the epidermal annulations, we have series of repeated parts of which the latter two

1 The choice of the unit is often a matter of convenience. 2 Apart of course from the primitive segmentation into

Thus in Text-fig. 121, p. 666, the unit of variation selected is _ the three regions, proboscis, collar, and trunk.
collar length x 4.

THE ENTEROPNEUSTA. 669
usually correspond fairly closely (cf. Willey, 799, Pl. XXVI. fig. 1, and Spengel, 703,
Pl. XXIV. fig. 8). With regard to the second part of the question it may be easily demon-
strated that in a given stretch of body length the branchiae are more numerous than are
the epidermal annulations or the hepatic caeca in a stretch of the same length in the
post-branchial region. The number of tongue bars averages about 5°6 per mm. of length
(see Table 14, p. 675). On the other hand the number of epidermal annulations in the
post-hepatic region is about 16 per mm. Therefore if we look upon these repeated parts
as evidences of a rudimentary segmentation’, we must regard the post-branchial segments
(or at least those of the hepatic caudal region) as being on the average 3°5 times as long
as the branchial segments. In other words the branchial segments are three times as
numerous as the hepatic-caudal ones per unit of length. Consequently if we calculate the
coefficients of variation on the basis of “segments” instead of collar length units we must
multiply the coefficient of variation for hepatic-caudal region by 3°5 in order to compare
it directly with that for the branchial region. This brings it up to the value 41:26 as
compared with 29°66 for the branchial region. If therefore we regard the Enteropneusts as
exhibiting a rudimentary form of segmentation the variability in the different body regions
falls into line with that of the Holochorda*. Much phylogenetic stress however must not be
laid upon this point since it is not improbable that the phenomenon of cephalization may
be shewn to be widespread among the Invertebrates also. What is of more importance is
the influence, small though it be, which the occurrence in the Enteropneusta of a form of
segmentation, similar in kind though less marked, may exert upon our conception of the
manner in which the merism of the Vertebrata may have arisen. And here a few remarks
of a more speculative nature may not be irrelevant in connection with

METAMERISM IN THE ENTEROPNEUSTA.

Willey 99, p. 303) has recently made a brilliant suggestion with regard to the origin
of gill clefts. The discovery of the peculiar dermal pits of Spengelia has led him to regard
gill-slits as having originally arisen as perforations in the interannular grooves formed for
the aeration of the gonads, a function which they eventually gave up for that of respiration
on the establishment of an elaborate vascular system. In this way Willey establishes a
connection between the segmentation of the gills and of the epidermal annulations, the
connecting link being the gonads. It has seemed to me possible to extend this conception
of Willey’s and to see in the gonads and their arrangement the prime factor in the
segmentation of the chordata. We may take as our starting-point a small Triploblastic
(and perhaps also Tricoelomate) creature with three more or less definite body regions, of
not widely unequal length, and of which the hindermost contains the gonads. The repro-
ductive elements would be small and fertilization external in the sea. Under these conditions
it is obvious that, other things being equal, those with the largest gonads would have the
best chance of leaving offspring to perpetuate and amplify this feature. Increased size of

1 Spengel (’03, p. 276 seq.) disagrees with Willey’s view
of the significance of the epidermal annulations and sees in
them only the effects of muscular contraction on the dis-
position of the epidermal glands. He points out that there
exists a certain amount of irregularity. Such irregularity
however seems to me in no way to militate against the
view of the significance of these structures taken by Willey.

Goer.

° It is interesting to notice that in Ptychodera the differ-
ence between variability in the different regions of the body
is not nearly so marked as in Amphiorus where we may
regard the cephalization process as having extended further.
Ihave found the following values for C.V. in different regions
of the latter genus. Preatrioporal segments, C.V.=1-792;
postatrioporal preanal, C.V.=2°655; postanal, C.V.=6-185.

86

670 R. C. PUNNETT.

gonad spells increased bulk and in the absence of a specialized vascular system this increase
of bulk must be almost entirely dependent on increased length. For respiration would depend
on transepidermal diffusion, a process which would more readily occur in a long slender
animal than in a short thick one. This great elongation of the gonad would necessitate
the establishment of accessory ducts by which its products could readily and rapidly escape.
From this it is but a short step to the stage of a series of independent gonads each with
its own duct extending throughout the elongated trunk region—a condition physiologically
comparable to that now found among the Nemerteans. Increase of bulk will still mean
increased fertility and such individuals will be favoured by Genetic Selection’, provided always
that the means for ensuring due aeration are adequate, whence we come to the establishment
of dermal pits for the aeration of the gonads. Later the pits become perforated and no
doubt these perforations supplied a physiological need and filtered off the excess of water
from the sand passing through the animal’s alimentary canal. Probably it was somewhere
near this phylogenetic epoch that the ancestral Enteropneusts took to an arenicolous life.
The problem of extraction of nutriment from a relatively enormous mass of innutritious
substance resulted in the specialization of the middle and hinder part of the digestive tract,
and the establishment of the hepatic caeca led to the disappearance of the gonads in this
region, They have left traces of their former presence in the epidermal annulations which
shew some tendency to irregularity now that their determining cause has disappeared; they
are still found in a rudimentary state in the hepatic region of some forms, but increased
physiological specialization has on the whole led to their confinement to the more anterior
portions of the trunk.

BIBLIOGRAPHY.
786. Bareson, W. “Development of Balanoglossus Kovalevskii,’ Quart. Journ. Mic. Se.
Vol. XXVI.
798. DELAGE AND HerRouarD. JT'raité de Zoologie Concrete, Vol. vi. Les Procordeés.

794. Hi, J. P. “On a new species of Enteropneusta from the coast of New South
Wales,” Proc. Linn. Soc. N.S.W. Vol. x.

798. Hu, J. P. “The Enteropneusta of Funafuti,” Memoirs Aust. Mus. 11.
785. Huprecut, A. A. W. Proeve eener ontwikkelingsgeschiedenis van Lineus obscurus. Utrecht.

702. Kuwano, H. “On a new Enteropneust from Misaki, Balanoglossus misakiensis, n. sp.”
Annot. Zool. Jap. Vol. Iv.

794. Morcaan, T. H. “The Development of Balanoglossus,’ Journ. Morph.

700. Rirrer, W. E. “Harrimania maculosa,’ Proc. Wash. Acad. Se. Vol. u.

702. Ruirrer, W. E. “The movements of the Enteropneusta,’ Biol. Bull. Vol. 111.

793. SpENGEL, J. W. “Die Enteropneusten,” Naples Monograph.

701. SpENGEL, J. W. “Die Benennung der Enteropneusten-Gattungen,’ Zool. Jahr. Vol. xv.
Syst. Abt.

708. SpENGEL, J. W. “Neue Beitriige zur Kenntniss der Enteropneusten,’ Zool, Jahr.
Vol. xvi. Anat. Abt.

799. Wiley, A. “Enteropneusta from the South Pacific, ete.” Willey’s Zoological Results,
Pt. 3.

1 Cf. Pearson, K., he Grammar of Science, 1900, p. 437, and also T’he Chances of Death, etc., 1897, Vol. 1. p. 63
(Essay on Reproductive Selection).

THE ENTEROPNEUSTA.

TABLE 11.

Pt. flava, var. laccadivensis.

671

5 eS Relative measurements a sii ees Relative measurements
a (in millimetres) (collar length=1) a (in millimetres) _ (collar length =1)
z aa = le =
piles pes (es) £ | $2 | sree) ore pees en epee ie
=
i |) 2 A == sls BESO 1 | 32,| 35] 8 = | oor) AS |)
2 hed 4 80 | 9-12 413° | 20:0 8 |) Be) |) Sp] == || eae 207 | —
Seed |) 4 ==) 1) 2:00, Heo Br 98} 35 | 74 | 250 | 383 | 246
Ao 3m 25 | 82 | 1-71 514 | 23:5 35 | 4 dee 11454200) an Waza es io
5 | 35) 35 | 75 | 214 | 500 | a1 36.~| 3 35 | 78 | 133 | 450 | 260° |
6 | 4 78 | 175 | 475 19°5 a7) 4 — | 225 | 4-75 egy
7 | 35 | 35 | — | 200 | 6-00 ese! 4 76 | 137 | 437 | 190
Breese a sh! | = Al es-00 =| 39 | 35 | 4 73 | 114 | 528 | 20:8
9 | 4 | 4 — || ies 5°50 | ay | 8 35 | 76 | 3:00 466 | 25:3
10 | 3 | 25 | 60 | 183 | 417 | 200 | 41 | 35 | 35 | 83 | 214 | 471 23:7
ie) | 35) | 35 || 90 || Sv i be4ae | e57— 4a! 4 4 ==) S219) 8) | bseo) |
Hee. | a5 | | ee + | 43 | 35 | 4 — | 200 | 5°85 ins
DBei|! 24 ay = POO a 44 | 4 4 a) SS | Ges —
14 | 35 | 35 | 105 | 175 | 443 | 300 || 45 | 35 | 4 == |) CRs Gal =
15 4 al a Siete | selon 46 | 3 $5) |) ese | 4-16 =
IG e4 | je 4 — | 300 | — | eyo! 3 — | 163 | 416 | —
leper ==) | Teg fl | 48 | 5 5 | 120 | 2:80 — | 240
Ise |e | 13 — | 300 | 466 | 49 | 5 5 | 110: ) 2:30 — | 22-0
1M) UN Ba ee reas iG ay 50.15 5 280 = =
20 | 25 | 25 | 60 | 180 | 420 | 240 | bl | 5 5 — | 200 = — |
at 3) | == |, 200. e600 |, 52 | 5 5 93 | 260 | 420 | 186
pp) 33 3 |e = 02,00 Ry | 8 5 2:00 = =
23 | 35] 35 | 74 | 185 | 5:27 54 |. 5 5 — | 2:20 skh Nee fa
SMBs) So |) = secagh || = 55 | 5 | 5 |140 | 270 | 350 | 280
95 || 4 ge (ee 300 Fn 56 | 5 5 | 125 | 2:50 BO). |) 25-09}
a / 3 | 3 | — | 266 | 467 Women |b 5 {125 | 220 | 540 | 250 |
Deke. [23 — | 200 | 466 58 OB 5 =| 50) )) 6s0 = |
Sl ay eae 35 | 137 | 437 | 212 || 59 | 45 | 45 | — | 333 | — =
a9 | 35 | 35 | 74 | 228 | 4:85 | 6 | 5 | 5 | — | a10 | 420
go | 4 | 4 | 93 | 295 | 3:50 32 || 61 | 5 | 5 | 11 | 190 | 610 | 222
Si eres eh Is. | 437 | 62 | 5 5 ‘| 105 | 2:20 | 3:80
} } I ml

Nos. 1—47 (inclusive) =Group 1.

”

=Group 2.

Nos. 93—119 (inclusive)=Group 3.

”

In Group 1 Collar length x breadth is not greater than 16.

» 48—92
” ” 2
” ” 3
” ” 4

”

”

”

” x
” x
»

”

”

is greater than 16, but not greater than 25.
is ” » 25, ” ” ” 36.

120—123 > =Group 4.

672

R. C. PUNNETT.

TABLE 11 (continued).

F mi’ Bie: ae Relative measurements a qe onelt ats Relative measurements
2 (in millimetres) (collar length = 1) E (in millimetres) (collar length=1)
3 %
a ao |e 2) a & os = a Sern eeuete lees 2 202 =
es |oslo sos E &2 | A ~2 |O2/og |r a ES gf iS
| |
63 | 45 | 45 | 9¢ | a1 | 477 | 2 oa |e || ey (aco: |) 350) -|! er
64 | 5 5 100 1-80 4-80 2 95 5 6 112 | 2:60 a 22-4
65 | 5 5 3:00 2:29 = 96 | 6 5 2) 9508.
66 | 5 5 2:20 4-40 _ 97 | 6 6 == || vag | = =
(Ge || cH |) 285 | — | Sem = 98 5 6 _ 2-20 ee ie
Ge || 2 |) & 110 | 166 | 5:89 99 | 6 6 =i || Sk) | = =
69 | 5 5 115 | 2-60 7:00 23-0 100 | 6 6 | 200 4:33 at
| G@ | 3 5 = 3:00 4:60 = 101 5 6 _ 3:40 _ =
a || 45 | — | 1:50 6:90 = 102 | 6 5 — | 2-00 seh
TON 5 5 00 = = 103 | 6 5 = |) 2:83 = a
| 73 |) & 45 | 108 | 2-00 4:60 216 || 104 | 55 | 55 | — | 255 — | =—
| A 5 105 | 2-00 5-60 21-0 105 | 6 6 =) |) 2:00 5:33
[Para ats aS — | 200 | 3:00 — |jwe | 6 | 5 | — | 166 = His
| 76 aes || oa te ery 6:00 a 107 5 6 | 85 1:60 260 | 170
Navas ied 5 = |) aso 4-60 _ 108 | 5 6 100 | 2-20 540 | 20:0
es || & 5 = | 2K) = | is 109 5 55 | 180 | 2:40 6:00 | 260
@ | 45 | 4 = | 135 Ba A || = iG) |) 6 109 | 2-40 440 | 218
80 | 45 | 45 | — | 1-44 =| = 111 5 Go| = 1:70 430 | —
81 5 4 = | 10 = = 112 5 55 | 136 | 2:80 480 | 27-2
82 | 5 en ee s76 B20) i a 113 | 6 6 140 | 1°91 4°75 23:3
83 | 5 5 — | 290 3:40 _ TGS 55 | 116 | 1:70 550 | 23-2
[B49 4! Be | A |) Se 512 19:0 115 | 5 55 | 123 | 2°50 4-70 24-6
8 | 5 45 | — | 1:50 4-10 _ 116 | 6 55 | 138 | 1:50 5:33 23-0
Sei la5 4-5 O) 4-20 = une lr GB SH | = | De 425 =
| ay || & 4 102 | 1:80 4:40 20-4 118 | 5 55 | 116 | 2°50 4:90 23:3
88 | 4 45 | 98 | 2:95 4°75 24-5 119 | 5 5:5) a ea:00 600 | —
89 | 5: 5 3:80 640 | — 120 | 7 75 | 140 | 2-28 3°71 20:0
90 | 5 5 — | 300 ESO! i 121 Gia || = | 38%) a =
91 5 5 = | 210 550 | — 122 | 6 i 145 | 3:16 _ 241
92 | 5 5 e240 380) (= WR || Uy 7 = | 260 ati =
Gan inom eee in= = eane eer
|

THE ENTEROPNEUSTA.

TABLE 12.

Pt. flava, var. muscula.

673

Pt. flava, var. gracilis.

g a pelle ee te | Relative measurements a | a fae | Relative measurements |
3 (in millimetres) | (GRIP eso 3 | (in millimetres) (collar length=1) |
a ee | s 2 | | ae ee E an = ee
my | oS |S 5 ehh is oa Ee eels Cee i] ts Sie claelh ba
| , | liars |
1 25 3 31 | 2:20 1:20 12-4 1 3 3 | 65 | 1-66 = | 21°6
2) | 43 4 48) || 2:33 ||, 3:66. 16009) een ae 2:5 | 68 | 2:00 Sa oO
omoae ie eb) 37 .| 250° |) -avspm ee iear allo ee 2 BT es | eo
4 2°5 3 35 2°20 3:40 14:0 4 2 25 | 56 2-000) |e wae 28-0
5 25 | 3 | 43 2°80 | 1:60 17:2 5 3 3 _ 183 |) — —
Gulica- | 35 | — |: 200 || 200 = Goleta oo Haas 1) edo, I, Sel Be
a 3 3d —_— 2:00 3°33 — 7 3 3 == | 1°33 \eaa =
8 By eS) — 2°50 1°83 - 8 2 2 == || isa) — =
9 25 35 — 2°40 3°80 — 9 2 2 =| 1:25 = | = |
TOME 2:5 4-3 74/0 200) Wiese SV HON a Leis! eb edie oie de33) iy) = rey |
a : Arist tlie rey fetch vrei} (yh =
|| 22 |) SR || 2) eg) |) eg = 19°6
| 13 2 2°5 _ 1:50 — —
14 2°5 2°5 40 | 1:20 — 16:0
15 | 2 2 36 | 1-00 _ 18:0
Pt. flava, var. parva. | 16 2 25 | 44 1°50 = 22-0
, } ; 17 2°5 3 44 1:20 _ 176
Relative length of branchial region (collar | oe ,
length=1) in 9 Specimens= ic SUA = 10 i%, x
1-20, 1:38, 138, 1-40, 1°60, 1-60, 1°60, 166,200. || 19 | 25 | 3 | — | 120 a a
|| 20 2°5 2°5 42 | 1:00 -- 16°8
Collar length in each case was 2°5-—3 mm. | 91 2:5 25 26 | 1-20 Ce 1 ea
In one complete Specimen total length | 22 2°5 2-5 42 | 1:00 2=.4'| fies
=30 mm. and relative length (collar length=1) | 23 3 2:5 44 1:00 A’ 146
was 12:0. 24 2 25 l= 1:25 — | =
| 25 | 25 | 25 | 40 | 1-00 == |) 16:0
| 26 2 2 32 1:00 — 16:0
|| 27 2 2 34 1-00 —_— 170
| 28 2 2 — 1:00 a ay
| | |

674 R. C. PUNNETT.

PAB ERs

Pt. flava, var. maldwensis.

|

g oa Daria eaeea | Relative measurements g i Fee Relative measurements
£ (in millimetres) | (collar length = 1) | & (in millimetres) (collar length=1)

eo vo

S eas |/e5|/e23 2s 3S r) es len | es s 2s 3
= | |

1 25 | 3 ss 1:40 3:60 — | a 4 4 = 175 3°75 =

B} |) ey |i By = 2:00 5:00 = 28 3 3 = 150 3:83 =

Sy) ee | 8} = 2-20 5-20 et: 29 | 3 3 = 1°50 4:50 _

4 | 3 25 | 48 1:00 4-66 160 || 30 | 3 Ba 6p 1:83 7-16 216

i | 3 |) 8 = 1:40 3:40 — 31 35 | 3 68 1°57 3:42 19°4

6) 225i 3 = 1:60 6:20 = 32 3 35 | 50 1°50 3°50 16°6

|| Ciel B 48 1:40 4:60 192 || 33 | 35°] 35 | 62 1-42 6:28 19-0

8 | 25 | 25 | 43 2:00 3°80 17-2 eyo] e 35 | 64 2°50 4°33 21°3

9 25 3 = 1:80 6°80 = 35 a) 3 — 1:14 SH
1@ || 25 |) 8 = 2-40 5-40 =e 36 | 3 3 | — 1:00 4:33 =
11 25 | 3 = 1:80 4-40 = 37 3 2 1:00 3:33 =
12 |.25 | 3 = i" 1:40 4:25 = 38 3 3 = 1:66 3-00 =
18) || 2H |) 8 = 2:00 4:60 = 39 | 3 3 = 1°66 3-00 =
14 | 25 | 25 | 41 1:80 4:60 16-4 40 | 3 3 38 1:16 3:33 12°6
i. || PH) B 39 1:40 3:80 156 || 41 35 | 4 = 1°71 5°85 =
16 | 3 25 | — 1:33 3°66 = ADS ES 3 = 1-50 4:16 _
17 2 25 | 29 1:00 2°75 145 ASS 3 S| SE 4:00 asi
18 | 25 | 3 = 1:60 5-00 oe ee a8} 3 41 1:16 3:83 13°6
19) 9-5) 4; 3 ==) || 1-60 5:40 = Ga fees Gs) — | 2-00 6°50 =
2a) IP SHB 1) BH || — 157 | 3-00 = 46 | 3 | 3 | _ 1:50 4-83 ae
21 3 35 | 72 3-00 5:00 24-0 as} yf | 4:16 =
a2 | 4 4 — | 137 | 312 — | 4 ee ie a ee (sie ceo es
23 | 3 3 = 1:33 2°33 — | 49) 4 45 | — | 3:50 625 Ee
24 | 4 4 80 1:87 412 20:0 | 50 | 4 45 | 68 1:37 4-50 16°5
25 | 3 3 =) || 22:16 4°16 _ 51 4 45 | 75 1:37 5°50 18°7
26 | 3 3 — | 200 | 333 e| 52 | 4 45 _ 2-25 5:37 =

| | | i

Nos. 1—19 (inclusive)=Group 1. Nos. 20—47 (inclusive)=Group 2. Nos. 48—52 (inclusive)=Group 3.

In Group 1 Collar length x breadth is less than 9.
3 Pe eX a is not less than 9, or greater than 16.
» i) Oe ip AY eine ap is greater than 16.

THE ENTEROPNEUSTA.

TABLE 14,

675

Pt. flava, var. laccadivensis.
| | |
| No. of Length Tongue bars No. of Length Tongue bars
tongue ee in mm. | per mm. | tongue bars | in mm. per mm.
— I | | =
104 10) he pea he 9°5 | 58
98 18 54 || 52 10 5:2
83 14 59 || 50 95 53
79 14 56 50 9°5 53
74 15 49 || 49 8 61
71 135 53 || 48 9 | 8
70 45 | 48 || 48 75 6-4
66 reed ese eae jee 8 59
64 10 64 || 40 ii 57
| 62 120s eet: 2a | 20 7 See
62 ul 56 || 38 6 63
59 10 59 34 55 62 |
57 10 57 || 32 5:5 58
Be ro 56 || 30 45 66
Average | 71:8 31 | 55 || 438 76 58

EXPLANATION OF PLATES.

advehent vessel of proboscis.
anterior neuropore,
basement membrane.
branchial pouch.

blood vessel.

collar coelom.

= collar canal.

= chondroid tissue.

central lumen of stomochord.
cornua of nuchal skeleton.

nuchal collar coelom (containing muscles).

connective tissue.

vascular ring of collar.

dorsal nerve.

dorsal proboscis coelom.

dorsal septum,

dorsal vessel.

external longitudinal muscles of collar.
efferent proboscis vessel.

= gonad.

ganglion cells.

ged.
gen.

glo.
gpl.
il.
int.
Ide.
7.
lm.
ls.
lu.
me.
mdv.
ml.
mle.
mld.
ml.
mlv.
n.

ngl.

dorsal ganglion cells of collar cord.
ventral ganglion cells of collar cord,
gland cells.

glomerulus.

Ist gill pouch diverticulum.

internal longitudinal muscles of collar.
intestine.

left diverticulum of dorsal proboscis coelom.
lateral lumen of stomochord.
longitudinal muscles.

lateral septum.

lumen of collar cord.

circular muscles,

dorso-ventral muscles.

longitudinal muscles.

longitudinal muscles of collar.
longitudinal dorsal muscles.
longitudinal lateral muscles.
longitudinal ventral muscles.

nuchal skeleton.

gland cells of collar cord.

676 R, C. PUNNETT.
nk. = keel of nuchal skeleton. per. =vight proboscis end vesicle.
nl. = layer of nerve fibrils (Punkt-substanz). vy. = either (1) racemose organ or (2) dorsal root.
0. = oesophagus. rde. = right diverticulum of dorsal proboscis coelom.
oep. = oesophageal epithelium. s. = stomochord.
07 = oesophageal nerve. Se. : : 3
abate : : NS cavity of lateral diverticulum of stomochord.
p. = proboscis coelomic cavity. sid.
pb. =parabranchial ridge. sk.1. = nuchal skeleton.
pbg. = post-branchial groove, skg. = branchial skeleton.
per. = pericardium. th, = tongue bar.
ph. = perihaemal space. v. = vermiform process of stomochord.
pg. = pygochord. em. = ventral nerve.
pig. = pigment. vpe. = ventral proboscis coelom.
pnp. = posterior neuropore. wv. = ventral recurrent vessel of proboscis,
pp. = proboscis pore. vs. = ventral septum.
pph. = peripharyngeal space. vv. = ventral vessel.
pvl. = left proboscis end vesicle.
PLATE XXXVII.
Fig. 1. Ptychodera asymmetrica, x 3.
Fic. 2. Pt. viridis, ventral view. Slightly enlarged.
Fic. 3. Balanoglossus carnosus. Slightly reduced.
Fig. 4. Pt. flava, var. gracilis. x 2.
Fic. 5. Pt. flava (var. maldivensis?). Regenerating. x 1.
Fic. 6. Pé. viridis, dorsal view. Slightly enlarged.
Fic. 7. Pt. viridis, lateral view. Slightly enlarged.
Fic. 8. Pt. flava (var. maldivensis?). Regenerating. x 1.
Fic, 9. Pt. asymmetrica. A large specimen. Slightly enlarged.
PLATE XXXVIII.
Fic. 10. Ptychodera flava, var. gracilis, Preserved specimen from Hulule, x 3.
Fie. 11. Pt. flava, var. parva. From preserved specimen, x 3.
Fic. 12. Pt. flava, var. cooperi. From preserved specimen. A small portion of the caudal end
probably missing. x 5.
Fic. 13. Spengelia porosa. A small portion of the hepatic region. x 4.
Fic. 14. Willeyia biswleata. Anterior end seen from dorsal surface. ~ 3.
Fic. 15. Balanoglossus parvulus. Section through the collar cord. A small clot of mucous substance

Fic.

is seen in the lumen. x 85.

16.
off the collar cord.

Pt. flava, var. parea.
x 20.

Section through anterior collar region with the Ist dorsal root coming

Fig.
Fic.
Fic.
Fic.

Fic.

Fic,
Fie.

Fic,
Fic.

Fic.

Fic.

Fic.

Fic.

Fic.

Fic.

Fic.

Fic.

Fic.

Fic.

Fic.

THE ENTEROPNEUSTA. 677

17. Pt. flava, var. saxicola. Dorsal view, slightly enlarged.
18. Balanoglossus parvulus. Anterior end from dorsal surface. x 6.
19. Pt. flava, var. gracilis. Preserved specimen from Minikoi. x 3.

20. Pt. flava, var. parva. Section through collar cord, The dorsal and ventral walls of the
central lumen shew a tendency to stick together. x 85.

21. Pt. flava, var. parva. Section similar to above but from another specimen. The central
lumen shews a marked tendency to occlusion and the lateral gland cells (g/.) are strongly
developed. x 85.

22. Balanoglossus parvulus. Section through proboscis pore. x 45.

23. Willeyia bisulcata. Section through collar cord. x 85.

PLATE XXXIX.

24. Balanoglossus parvulus. Section through collar. x 45.

25. Pt. flava, var. laccadivensis. Section through proboscis. x 45.

26. Pt. flava, var. parva. Section through anterior collar region. x 20.

27. Pt. flava, var. maldivensis. Section through edge of left genital pleura, x 85.

28. Pt. flava, var. saxicola. Unusual dorsal root formed by pigmented outgrowth from collar
cord and glandular ingrowth from epidermis. These two portions fuse a few sections further
back. x 85.

29. Willeyia bisulcata. Section through proboscis towards its hinder end. The dorso-ventral
muscles (mdv.) are seen converging to the minute vermiform process of the stomochord. x 45.

30. Pt. flava, var. saxicola. Small portion of inner surface of genital pleura shewing ingrowth
from epidermis forming gonad, x 110.

31. Pt. flava, var. saxicola. Section through tip of a genital pleura shewing epidermal ingrowth
forming gonad. x 180.

32. Pt. viridis. Section through proboscis, x 20.

PLATE XL.

33. Pt. flava, var. parva. An immature gonad shewing ectodermal connection, and quantity
of mucus inside. x 180.

34. Spengelia maldivensis. Section through collar canal just anterior to its opening into the
first gill pouch, Ventral to it is the blind diverticulum of the first gill pouch of which the
internal opening is marked by an asterisk *. x 85.

35. Pt. flava, var. parva. An immature gonad in which the ectodermal connection is occluded
by a mucous plug. x 180.

36. Pt. flava, var. laccadivensis. From young specimen (anterior hepatic region) shewing very
young gonad connected with the ectoderm. x 410.

G.I 87

678

Fic.

Fic.

Fic.

Fic.

Fic.

Fic.

Fic.
Fic.
Fic.
Fic.

Fie.

Fig.

Fic.
Fic.

Fic.
Fie.

Fic,
Fie.
Fie.
Fie.
Fie.
Fic.

R, C. PUNNETT.

37. Pt. flava, var. muscula, A nearly ripe ¢ gonad breaking through to the ectoderm.
x 180. (sp=spermatocytes.) From an iron haematoxylin preparation.

38. Pt. flava, var. laceadivensis. From young specimen. a—e represent stages in the formation
of the giant nutritive cells. All x 520.

39. Pt. flava, var. laccadivensis. Young gonad, shewing cavity formed inside, and outer coat
of nutritive cells forming a kind of follicle. The membrane surrounding the gonad derived
from the lateral septum is still intact. x 410.

40. Pt. flava, var. laccadivensis. Young gonad, slightly older than in preceding figure. The
membrane surrounding the gonad tends to break down in places. «x 410.

41. Pt. flava, var. saxicola. Section through young gonad with ectodermal connection. At the
base of the gonad the membrane is deficient permitting the invasion of nutritive cells. x 180.

42. Pt, flava, var. saxicola. Gonads somewhat older than in preceding figure. x 110.

PLATE XLI.
1. Balanoglossus parvulus. Section through branchial region. x 45.
2. B. parvulus. Through region of post-branchial groove. x 45.
3. B. parvulus. Through nuchal skeleton in anterior region of collar. x 45,
4. B. parvulus. Through post-branchial genital region. x 45,
5

Spengelia porosa, Through hepatic region. On the right the section has gone through
the middle of a hepatic caecum. x 12.

6. Sp. maldivensis, Section through nuchal skeleton in the region of the keel (nk.). The
perihaemal cavity (ph.) of the right side is seen at this level. That of the left side does
not reach so far forwards. x 45.

7. Sp. porosa. Section through the intestinal region behind the hepatic caeca. x 12.

8. Sp. maldivensis. Schematic longitudinal sagittal section reconstructed from transverse
sections. x 10.

9. Willeyia bisuleata. Section through proboscis pore. x 45.

10. W. biswleata. Schematic reconstruction as in fig. 8. x 10.

PLATE XLII.

ll. Spengelia porosa. Section through posterior part of proboscis. x 12.
12. Willeyia bisuleata. Section through anterior part of proboscis. x 30.
13. W. bisuleata. Section through branchial region. x 22.

14. W. bisuleata. Section through genital region. x 22.

15. Wz. bisulcata. Section through hinder part of anterior neuropore. ~ 45.

16. W. bisuleata. Through anterior part of anterior neuropore, shewing. the cesophageal nerve
coming off on the left side of the figure. x 45.

THE ENTEROPNEUSTA. 679

Fie. 17. Ptychodera viridis. Section through post-branchial groove. x 12.
Fic. 18. Pt. flava, var. muscula. Section just anterior to the proboscis pores, shewing the expansion
of the nuchal skeleton over the racemose organ (7). x 45.

Fie, 19. Pt. viridis. Section through anterior collar region. x 20.

Fie, 20. Sp. maldivensis, Section through post-branchial groove. x 45.

PLATE XLIII.
All figures on this plate are from P¢. flava, var. parva.

Fig. 21. Through anterior part of proboscis. x 30.
Fic. 22. Through anterior portion of central proboscis complex. 30.

Fic. 23. Through specimen in which the two proboscis pores have a common opening near the
mid-dorsal line. x 54.

Fig, 24. Through hinder part of proboscis shortly before the lateral diverticula of the stomochord
appear. x 54.

Fie. 25. Through region of the lateral diverticula of the stomochord. x 54.

Fie. 26. Through the proboscis pores behind the lateral diverticula of the stomochord. x 54.
Fie. 27. Through anterior cavity containing region of collar. x 30.

Fic.

to
ie)

Through region of racemose organ, behind the proboscis pores. x 54.

PLATE XLIV.

Fie. 29. Pt. flava, var. asymmetrica. Section through collar cord. x 45.

Fie. 30. Pt. flava, var. parva. Section through same level as Pl. XLIII. fig. 25, but through
another specimen in which the lumen of the stomochord is obliterated instead of being patent.
x 45,

Fie. 31. Pt. flava, var. parva. Section through middle of collar region shewing longitudinal muscles
arranged in bundles. x 20.

Fic. 32. Pt. flava, var. parva. Section through proboscis pores, In this specimen the perihaemal
spaces reach forwards to this level. More usually they commence shortly behind the pores
(cf. Pl. XIII. figs. 23, 26). . x 45.

Fie. 33. Pt. flava, var. saxicola. Section just anterior to the proboscis pores shewing great width
of keel of nuchal skeleton. x 45.

Fic. 34. Pt. flava, var. saxicola. Section through stomochord in region of lateral diverticula. x 85.

Fies. 35, 36, 37. Pt. flava, var. saxicola. Sections through the pygochord. The three sections are
from the same specimen; fig. 35 is nearest to the anus, fig. 37 is furthest from it. x 85.

Fie. 38. Pt. flava, var. saxicola. Through anterior branchial region, x 12.

Fic. 39. Pt. flava, var. saxicola. Through posterior branchial region of same specimen as in fig. 38.
x 12.

Fic. 40. Pt. flava, var. saxicola. Through level immediately behind the post-branchial groove. Also
from same specimen as fig. 38. x 12.

Fie. 41. Pt. flava, var. sawicola. Through region of post-branchial groove. The genital pleurae
are still almost as large as in the branchial region. «x 12.

87—2

680 R. C. PUNNETT.

PLATE XLV.
Fic. 42. Pt. flava, var. gracilis. Section through hinder part of collar region. «x 45.

Fie, 43. Pt. flava, var. laccadivensis. Sections through stomochord. All x 85.
a. Near anterior extremity.
b, At anterior level of lateral diverticula.
c. Through lateral diverticula at their maximum breadth.

d. In region of nuchal skeleton.
Fie. 44. Pt. flava, var. gracilis. Through stomochord, shewing completely obliterated lumen. x 85.
Fic. 45. Pt. flava, var. cooperi. Through keel of nuchal skeleton. x 85.
Fie. 46. Pt. flava, var. cooperi. Through proboscis stalk close to opening of proboscis pores. x 85.

Fic. 47. Pt. flava, var. cooperi. Through post-branchial groove. The groove is large and the
genital pleurae well marked. «x 45.

PLATE XLVI.

Sections through the region of the branchial and post-branchial groove in several species and
varieties of Ptychodera. All x 27.

Fic. 48. Pt. flava, var. parva. Through middle of branchial region.

Fic. 49. Pt. flava, var. laccadivensis. Through post-branchial groove.

Fic. 50. Pt. flava, var. gracilis (Hulule). Through middle of branchial region.

Fie. 51. Pt. flava, var. laccadivensis. Through hind end of post-branchial groove.
Fic. 52. Pt. asymmetrica. Through middle of branchial region.

Fie. 53. Pt. flava, var. gracilis (Hulule). Through hind end of post-branchial groove.
Fie. 54. Pt. flava, var. gracilis (Minikoi). Through branchial region.

Fig. 55. Pt. flava, var. gracilis (Minikoi). Through hind end of post-branchial groove.
Fig. 56. Pt. asymmetrica. Through posterior branchial region.

Fie, 57. Pt. flava, var. laccadivensis. Through middle of branchial region.

Fie. 58. Pt. asymmetrica. Through hind end of post-branchial groove.

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MARINE CRUSTACEANS.
PARTS X. AND XI

By L. A. BorrapatLe, M.A., Lecturer in Natural Sciences at Selwyn
College, Cambridge.

(With Plates XLVII. and XLVIII. and Text-figures 122—6.)

X. THE SPIDER-CRABS (OXYRHYNCHA).

THE Oxyrhyncha are a fairly well characterized group. Sharply separated, on the one
hand from the Dromiacea by the coxal opening of the oviduct, the loss of the first
abdominal limb in the female, and the normal shape and position of the last pair of legs
(except in two or three cases as Ocinopus (Fig. 123), Grypachaeus and Zebrida), and on
the other hand from the Oxystomata by the square mouth and the normal maxillipeds
(Pl. XLVII. fig. 5b), they differ from the Brachyrhyncha in the shape of the body, which
is narrowed in the fore part, so that in most cases it is triangular, and has a well-marked
rostrum (Pl. XLVII., and Text-figs. 122 and 123), and usually in the imperfection of the orbits.
Slender legs and weak chelae are another common feature, but not a diagnostic one. Their
prevailing habitat may be summed up by saying that they are the crabs of the weed and
weed-like animals, but this statement needs some explanation and qualification.

The loss of their tail-fin has left the crabs, as a whole, a distinctly slower-moving
group than most of the tailed Reptantia. To this there are, indeed, exceptions—as, for
instance, the Ocypodes on land and the swimming crabs in the water—but for the bulk
of the Brachyura it holds good, and each of the sections of the tribe has had to meet
the difficulty in its own way. In the Brachyrhyncha the cuticle has generally become
thickened into a stout armour, while the crab keeps fairly active in its habits, at all events
after dusk, and usually defends itself vigorously with its strong chelae when it is attacked.
In this legion protective coloration and shape are comparatively rare, and the habit of
covering the body with foreign objects decidedly so (Caphyra), though the custom of hiding
under stones is common. The remaining groups have, so to say, accepted the situation.
Driven by the loss of their tail-fin to lead a sluggish life, they have found safety in
exaggerating this inertness and combining it with various devices for escaping observation.
The Dromiacea hold foreign bodies over their backs with their hind legs, the Oxystomes
bury themselves in sand or shingle, and in each of these groups the most characteristic
features, to which they owe their distinctness, are due to the method of concealment’. The

1 See above, p. 434,

682 L. A. BORRADAILE.

typical members of the Oxyrhyncha—the Maiidae'—have adopted another plan. They hide
themselves by covering their bodies with bits of seaweeds, zoophytes, or sponges, which are
held on by hooked or jagged hairs of special shape found on the body and limbs of the
true spider-crabs and not elsewhere (Pl. XLVII. figs. 3c and 4d). They are gathered and
placed in position by the crab itself, which uses for this purpose its chelipeds. These limbs
are specially shaped so as to have a mobility not found in other crabs, and are thus able
to reach distant parts of the body and legs. In order that they may adhere better, the
fragments of weed, etc. are treated with a secretion given out by glands on the first
maxilliped?; and they not only remain in a living state, but often continue to grow, so
that in some genera it is not uncommon to find the whole crab as completely hidden by
a single sponge as any Dromia. Besides seaweeds, the organisms used include sponges,
hydroids, polyzoa, and ascidians, and in some cases barnacles and tubicolous worms add
themselves as self-invited guests. Several different kinds of organisms may sometimes be
found on the back of one crab, but in most cases, very possibly in all, the species planted
by it are those amongst which it is living and are changed if it be placed in other
surroundings where they would be conspicuous. This implies considerable care in the choice
of the clothing, and indeed such may easily be seen to be bestowed if a captive individual,
say a Maia squinado, be watched while it is disguising itself. I have even seen this species
cover its back with shingle when no weed was available. The number of hairs naturally
varies, as does also the extent to which the crab is hidden, but it rarely happens (pzaltus)
that they are quite wanting.

1 Key to the families of the Oxyrhyncha. Second joint of antenna truncate-triangular. Hye-
I. Carapace thin and flat. First leg (cheliped) not long or stalks very short. Acanthonychinae.
specially mobile or with fingers bent at an angle with the B. True orbits, containing both supra- and postocular
hand. Male opening sternal. [No orbits. Second joint elements sheltering the eyes, are more or less com-
of antennal stalk slender, fused with epistome but not pletely formed, except in a few genera where the
with front. No hooked hairs. ] Hymenosomidae. eyestalks are long and slender. Second antenna-joint
II. Carapace not thin and flat (except Ocinopus). First leg broad, usually not truncate-triangular. Hyestalks long
either mobile or powerful with bent fingers. Male opening or short.
coxal. 1. A large, cupped, usually blunt postocular process
A. Chelipeds specially mobile, rarely much larger than present. Eyestalks short. Cornea of eyes not com-
the other legs or with fingers bent at an angle on the pletely hidden when they are folded back. Pisinae.
hand. Second joint of antenna well developed, generally 2. Postocular process, if present, usually sharp and not
fused with epistome and often with front. Orbits eupped, but if not so then cornea hidden (as also
generally more or less incomplete. Hooked hairs in most other cases). Eyestalks usually long.
almost always present. Maiidae. Matinae.
B. Chelipeds not specially mobile, usually much longer
and heavier than the other legs and with fingers bent Key to the subfamilies of the Parthenopidae.
on the hand at an angle towards the side on which I. Carapace usually triangular, sometimes suboval or sub-
the fixed finger is set. Second joint of antenna small, pentagonal. Rostrum simple. Chelipeds much bigger
short, and not fused with epistome or front. Orbits than the other legs. Branchial regions of the body
well made. Hooked hairs almost always wanting. deeply separated from cardiac, Parthenopinae.
Parthenopidae. II. Carapace usually sharply pentagonal. Rostrum cleft

Key to the subfamilies of the Maiidae. into ne june pad ee Ree

I. Second joint of antenna very slender throughout its length. cet i aa tas erase ohh ganze att oo

[No crete Eyestalks generally long.] COE: 2 For an account of those structures in the Spider-Crabs

II. Second joint of antenna not very slender. which are specially adapted to their habit of clothing them-

A. No true orbits (eyestalks hidden under a supraocular selves, see Aurivillius, Kong. Svenska Vet.-Ak. Hand. Bt.
spine or sunken in the sides of a great rostrum). xxu1. no. 4 (1889),

MARINE CRUSTACEANS. 683

Such habits as these need a corresponding habitat, and the Spider-Crabs are essentially
haunters of weeds, and weed-like animals. The larger kinds cling to the rocks and stones
on which grow the organisms they clothe themselves with. The walking-legs of such species
usually end in strong, sharp, curved claws by which they can hold fast to the ground.
Smaller kinds often live on the weed itself, and these frequently bear hooked or even
sub-chelate claws on the hinder legs with which to cling to the branches, while one or
both of the second and third pairs are long—probably because they are used in climbing
(Pl. XLVII. figs. 1, 3 and 4). The first pair (chelipeds) are also sometimes used, monkey-
wise, in clambering. Research will no doubt show that some of the peculiar features of
particular species and genera are adaptations to special kinds of sessile organisms. At present
I can only recall the flat, leaf-like body of Huenia, which resembles the Halimeda-weed
among which it is generally found (see below, p. 686), and it is certainly the case that
very many species show no preference whatever in this respect.

The habit of living on or among sessile organisms is probably kept up by some of
the members of the other two families of the Oxyrhyncha, although it is the Matidae
alone which clothe themselves in the way described above. Further information, however,
is much needed on this point. The Hymenosomidae (Fig. 122) seem clearly adapted by
their structure—their delicate bodies and slender legs with hooked end-joints—for living on
plants or zoophytes. They may certainly sometimes be found in such situations, but are
also reported to have been taken under stones!, a position for which their flat backs are
not unsuitable. Among the Parthenopidae, the Eumedoninae are probably guests of other
organisms. Zebrida has, indeed, been taken among the spines of a sea-urchin whose colouring
it assumes. But the Parthenopinae (Pl. XLVII. fig. 5) have an entirely different habitat.
The members of this subfamily have left the weed (that they originally had the same
habits as the Matidae seems likely from the occurrence of hooked hairs, slight and few in
number, in certain members of the genus ZLambrus) and have taken up the same _ habitat
as the Oxystomata—that is to say beds of sand and shingle. The result is a series of
modifications strikingly like some of those which are found in the latter group» The
overlapping wings of the carapace which hide the legs of Calappa and Tlos reappear in
such forms as Cryptopodia, Heterocrypta and Oethra and less strongly marked in Lambrus,
especially L. calappoides. The long chelipeds of some Leucosiids are repeated in an altered
shape in various species of Lambrus and Parthenope. The bent fingers of Ranina and some
species of Calappa are found throughout the group. In Lambrus calappoides we find again
the flat hands of Calappa held against the breast. But the most striking of these likenesses
is the arrangement of the inward channel for the breathing-stream in Aulacolambrus, which,
while it is wholly new, yet strongly recalls that of the Leucosiidae. In this subgenus, the
underside of the carapace (pterygostome) is traversed, on each side, just outside the third
maxilliped, by a deep groove which is covered in, not like the analogous channel in the
Leucosiidae by the maxillipeds, but by thick fringes of hairs borne by the maxillipeds and
by the carapace, becoming thus a closed tube leading from the front of the body to the
opening of the gill-chamber at the base of the chelipeds (Pl. XLVII. fig. 6). A similar

1 Adams and White, Crustacea of the Samarang. The ganisms. Or, again, it may be that the hollows under the
authors make the same statement about Trapezia, which is stones from which the Hlamena was taken were lined, as
certainly a coral crab. Perhaps these genera take shelter such hollows often are, with a scrubby growth of weed.
under stones when they are by some accident removed from 2 See above, pp. 434, 435.
the neighbourhood of their natural habitat, growing or-

684 L. A. BORRADAILE.

arrangement is found in the other subgenera of Lambrus'. In this case, however, there
are two shallow grooves, an outer one running outward and an inner one running forward
from the gill-opening, and both uncovered (PI. XLVII. fig. 56). The rough, knobbed back
of the Parthenopinae is like that of Calappa, but a similar texture is so common among
crabs that little can be argued from it. Probably it is less conspicuous on a sandy bottom
with coral pebbles than a quite smooth object would be.

We have already stated that the Oxyrhyncha are among the sluggish groups of crabs.
When they are seized they do not attempt to defend themselves with their chelae, but
move their legs feebly and aimlessly. Indeed the Parthenopinae—like the Calappidae in
this respect also—often seem to be hypnotised by being moved, drawing up their legs under
them and remaining still. In general the intelligence of the group is of a low order, with
the single exception of their cleverness in disguise, which, after all, is probably no more
than a fairly simple reflex.

Fic. 122. Elamena gracilis; a. whole animal, b. outside of hand.

Of the 29 species in the collection made by the expedition, three are new. The rest
are all recorded from the Indian region by Major Alcock?, with the exception of two which
were described as new by Miss Rathbun in her account of Prof. Agassiz’s Maldive Crabs’.
The following systematic list sets forth these species in order.

Family Hymenosomidae.
Genus Hlamena H. M.-Edw., 1837.

1. Hlamena gracilis, n. sp. (Fig. 122)
Diagnosis: “An Elamena in which the sides of the hinder part of the body are straight
from the 4th to the Ist pair of legs, and then turn inwards almost at right angles to

1 Owing to lack of material Iam unable to say whether Journ. As. Soc. Bengal.
this groove is found throughout Parthenopinae. 3 Bull. Mus. Harvard, xxx1x. 5 (1902).
2 In his series of papers on the Indian Crabs in the

MARINE CRUSTACEANS. 685

join the fore part, which is triangular, with very slightly irregular sides; the front also
triangular but with indications of two teeth at its sides; the eyes showing a small part
of the cornea at the sides of the front when seen from above; the chelipeds short, slender,
with narrow, spooned fingers about as long as the palms; and the walking-legs slender,
with a spine at the end of the meropodite, and the last joint strongly hooked, with a
group of little thorns underneath at the tip and a fringe of hair all along.”

Length of longest specimen: 6mm. Breadth: 7mm. Colour: in spirit, pale yellow;
when alive, legs black-brown, rest of body transparent.

A female was taken at Minikoi, and a male and female at Hulule, Male Atoll, all on
the reef.

Family Maiidae.
Subfamily Inachinae. Genus Achaeus Leach, 1815.

2. ?Achaeus spinosus Miers, 1879. Alcock, 1. p. 171’.

My specimen agrees with Alceock’s and Miers’ descriptions, but the last two pairs of
legs have strongly curved end-joints. Miers and Alcock only mention the last pair.

Dredged in Mahlos Atoll in 24 fathoms.

< frac
LASS
RSS

SS

Fic. 123. Ocinopus aranea; a. whole animal, b. two of the hairs on the hinder pair of legs, c. one of the long hairs
on the legs of the second and third pairs, d. hooked hairs.

Genus Ocinopus de Haan, 1837.
3. Ocinopus aranea de Haan, 1837. Alcock, 1. p. 183 (Fig. 123).

The shape of the lobes at the tip of the rostrum in this species varies somewhat.

A very common crab, taken practically throughout the group in 19—45 fathoms.

1 For a statement of the principles on which references Alcock’s paper on the Indian Oxyrhyncha appeared in Journ.
are given in this series of papers see above, p. 192. Major As. Soc. Bengal, uxtv. ii. 2, p. 157 (1895).

G. IL. 88

686 L. A. BORRADAILE.

Subfamily Acanthonychinae. Genus Xenocarcinus White, 1847.

4, Xenocarcinus tuberculatus White, 1847. Alcock, 1. p. 192.
Dredged outside Fadifolu Atoll, in 70 fathoms.

Genus Menaethius H. M.-Edw., 1834.

5. Menaethius monoceros (Latr.), 1825. Alcock, 1. p. 197.

A common species, taken in Male, Fadifolu, Minikoi, and Goifurfehendu Atolls down to
6 fathoms.

Genus Huenia de Haan, 1837.
6. Huenia proteus de Haan, 1837. Alcock, 1. p. 195 (Pl. XLVII. figs. 1, 2).

In the extraordinarily variable shape of its body, this crab is fully worthy of its name.
The simplest form is found in most of the males, which are flat and triangular, much like
Menaethius, with a long, sharp rostrum (Pl. XLVII. fig. 1).
What may be the habitat of these males is yet to be settled: .
the specimen figured carried a large flat piece of a green weed, SO
held on its rostrum by means of the hooked hairs always found ee
there in this species. On the other hand, most of the females
and some of the males—true males, not “unsexed females” with
parasites—have the body widened into a leaf-like shape by
outgrowths of the hepatic and branchial regions (Pl. XLVII.
fig. 2). These individuals, at least in most cases, live on the
flat Halimeda-weed, which they closely resemble both in shape
and colour. Figure 124 is given to show this lkeness. Some of
the females, however, come near the three-cornered shape of the
ordinary male. This, and the fact that intermediates are found
in both sexes, prevents us from callmg the phenomenon
“dimorphism.” Another interesting feature is shown by the
walking legs. The last joint in these limbs is strongly toothed,
and can be shut back at a sharp angle on the one before it.
In the broad individuals the legs are short and stout and
keeled to look like the edges of Halimeda “leaves,” and here

Fie. 124. A sprig of Halimeda-weed

i 5 = with a specimen of Huenia proteus,
the whole under-edge of the last joint but one is hairy; but showing the likeness of the latter

in the triangular form, where the legs are long and slender, toa “leaf” of the weed.

there is a special tuft of hairs to meet the end-joint, and this

tuft is often raised on a knob. The whole structure thus formed is all but subchelate and
seems clearly adapted for holding on by.

The surface of the body of these crabs may be seen under a high magnification to
have the curious graving shown in fig. 1a, on Plate XLVI.

The species was taken in Male Atoll on the reef, and dredged from 22 fathoms in
Kolumadulu Atoll.

MARINE CRUSTACEANS. 687

Subfamily Pisinae. Genus Nazioides A. M.-Edw., 1865.

The synonymy of this genus is discussed by Rathbun, Proc. Biol. Soc. Washington, XI.
p. 157 (1897).

7. Nasioides hirta A. M.-Edw., 1865.

Nawia hirta, Alcock, loc. cit. p. 218.

Taken in Kolumadulu, Suvadiva, Haddumati, and South Nilandu Atolls in 30—40 fathoms.

8. Nawioides spinigera, n. sp. (Pl. XLVII. fig. 3).

Diagnosis: “A Nasioides with the horns more than half the length of the body, bearing
the accessory thorn a third of their length from the end; the body hairy, with many knobs
and thorns, of which the largest are three in a row on the gastric region, two on the
cardiac, one on the intestinal, three in a triangle on the branchial, with one in front of
them, and one on each eye-hood, these latter being quite upright, but bent slightly forwards
at the tip, no thorn at the angle of the mouth, two knobs on the pterygostome, two
thorns on the outside of the basal joint of the antenna; the chelipeds slender, little longer
than the body, the fingers half the length of the palm, finely toothed and meeting along
nearly their whole length, a spine at the end of the arm; and the walking legs slender,
the first pair very long, a long spine at the end of the meropodite of this limb, a short
one on the next, and none on the hinder two.”

Length: 11 mm. Breadth: 8mm. Colour in spirit: white.
Allied to NV. cerastes (Ortm.), 1894.
Three males dredged in 30 fathoms in North Male Atoll.

Genus Halimus Latr., 1829.

For the synonymy of this genus, which is identical with Hyastenus White, 1847, and
not Nawia Leach, 1828, see Rathbun, Proc. Biol. Soc. Washington, x1. p. 157 (1897).

9. Halimus tenuicornis (Pocock), 1890. Alcock, 1. p. 215.

The colour of this species as preserved in formalin is white with bright pink markings.
It was dredged in South Nilandu, Male, Mulaku, Felidu and North Male Atolls in 25—30
fathoms.

10. Halimus gracilirostris (Miers), 1879. Alcock, 1. p. 215.

Dredged throughout the group in 20—70 fathoms.

11. Halimus diacanthus (de Haan), 1837. Alcock, 1. p. 210.
Found on a black crinoid dredged in 30 fathoms in South Nilandu Atoll.

12. Halimus calvarius (Alc.), 1895. Alcock, 1. p. 213.

Dredged in South Nilandu, Mulaku, Suvadiva, Haddumati, and North Male Atolls in
19—40 fathoms.

13. Halimus convexus (Miers), 1884. Alcock, I. p. 216.

The one specimen in the collection is a female, and differs from Miers’ (a male) in that:
(i) the horns are less strongly divergent, (11) the palm is slenderer and the fingers much less

88—2

688 L. A. BORRADAILE.

gaping. The species is allied to H. calvarius but differs in that: (i) the horns are more
slender and diverge more strongly, curving, as well as slanting, outwards, (ii) there is no
trace of a knob on the gastric mound, which is also more prominent, (iil) there is no
epibranchial spinule or intestinal spine, (iv) the carapace is narrower, and (v) the palm is
rather more swollen. It was dredged in 30 fathoms in Suvadiva Atoll.

14. Halimus espinosus, n. sp. (Pl. XLVII. fig. 4).

The group of forms akin to H. diacanthus (including var. elongata Ortm., 1893, H.
subinermis Zehnter, 1894, H. convecus Miers, 1884, and H. calvarius Alc., 1895) are in all
likelihood no more than varieties of one variable species. The collection contains a specimen
of another such form, which I describe here. The following characters, taken together, separate
it from its allies mentioned above: (1) carapace narrow (10:6); (41) gastric region swollen,
with traces of the knob; (iii) horns rather less than half the length of the carapace, straight,
not very divergent; (iv) a spinule on each branchial region and a low knob in place of the
intestinal spine; (v) the outer angle of the basal antennal joint not strongly marked. The
specimens, which are both males, were dredged in Haddumati Atoll in 35 and 39 fathoms.

15. Halimus agassizi Rathb., 1902.

Halimus agassizii, Rathbun, Bull. Mus. Harvard, xxxrx. 5, p. 138, fig. 6.

The females of this species have very slender chelipeds with long narrow palms and
short fingers, which gape very slightly. Miss Rathbun’s only specimen was taken in the
same locality as ours, Nilandu Atoll, and in practically the same depth, 24 fathoms.

Genus Phalangipus Latr., 1825.

Miss Rathbun (Proc. Biol. Soc. Washington, X1. p. 159, 1897) has shown that this name
must be substituted for Hgeria Leach, 1815, which is preoccupied.

16. Phalangipus arachnoides Latr., 1825. Alcock, 1. p. 223.
Dredged in Mulaku, South Nilandu, and Felidu Atolls, in 25—36 fathoms.

Genus Tylocarcinus Miers, 1879.

17. Tylocarcinus styx (Hbst.), 1803. Alcock, 1. p. 235.
Taken on the reef in Male, Fadifolu, and Goifurfehendu Atolls.

Subfamily Maiinae. Genus Schizophrys White, 1848.

18. Schizophrys aspera (A. M.-Edw.), 1834. Alcock, 1. p. 243.

Taken on the reef in Male and Fadifolu Atolls, and dredged in 34 fathoms in Felidu
and 20 fathoms in Suvadiva Atoll.

Genus Cyclax Dana, 1852.

19. Cyclax (Cyclomaia) suborbicularis (Stimps.), 1857. Alcock, I. p. 245.
Taken on the reef at Hulule, Male Atoll.

MARINE CRUSTACEANS. 689

Genus Micippa Leach, 1816.

20. Micippa philyra (Hbst.), 1803. Alcock, 1. p. 249.
Taken on the reef at Hulule, Male Atoll, and dredged in Kolumadulu, South Nilandu,
Fadifolu, Suvadiva and North Male Atolls in 20—35 fathoms.

21. Micippa margaritifera (Hend.), 1893. Alcock, 1. p. 253.
Dredged in Suvadiva Atoll in 43 fathoms.

22. Micippa parca Alc., 1895. Alcock, 1. p. 253.

I think that this form should have specific rank, rather than that of a variety of
M. margaritifera, given to it by Alcock.

Dredged in 20 and 25 fathoms in Mahlos and South Nilandu Atolls respectively.

Genus Macrocoeloma Miers, 1879.

23. Macrocoeloma nummifer Alc., 1895. Alcock, 1. p. 255.

Almost all the spines on my specimens end in a round knob. Alcock does not mention
this, but such knobs were present on several of the spines in the individual he figures.
In my specimen thickenings of a like kind are set on the tips of the front. Alcock’s
figure, however, shows none. The knobs cannot be rubbed off and do not seem to be due

to foreign growths of any kind.
The species was dredged in 30 fathoms in South Nilandu, and in 23 fathoms in
Mahlos Atoll.

Family Parthenopidae.

Subfamily Parthenopinae. Genus Lambrus Leach, 1815.

24. Lambrus (Rhinolambrus) bispinosus Rathb., 1902.

Lambrus (Rhinolambrus) bispinosus, Rathbun, Bull. Mus. Harvard, xxx1x. 5, p. 134,
figs. 1, 2.

In her description of this species, Miss Rathbun says that the upper side of the hand
bears two lobes. Later she says that the species is separated from JZ. confragosus (among
other differences) by the presence of only one lobe in this position. Her figure and my
specimens bear out the latter statement.

Dredged in South Nilandu Atoll in 25 fathoms.

25. Lambrus (Rhinolambrus) pelagicus Riipp., 1830. Alcock, 1. p. 267.

The granules on the back are quite well marked in my specimens, as in Milne-Edwards’
figure (Now. Arch. Mus. vit. Pl. XIV. fig. 4). It would seem that Alcock had found in
the Andamans local races both of this species and of ZL. gracilis, im which the granules
were reduced in size and distinctness.

Dredged in Mulaku, Suvadiva, and Addu Atolls, in 28—43 fathoms.

690 L. A. BORRADAILE.

26. Lambrus (Rhinolambrus) gracilis Dana, 1852. Alcock, 1. p. 269.

The specimens which I assign to this species agree with Alcock’s description fully,
except that the surface of the carapace is not absolutely smooth, there being several granules
round the cardiac spine and along the branchial ridges.

Dredged in Felidu, Fadifolu and Suvadiva Atolls, in 22—43 fathoms.

27. Lambrus (Rhinolambrus) turriger Ad. and Wh., 1847, Alcock, 1. p. 269 (Pl. XLVII.
fig. 5).

Dredged in South Nilandu, Felidu, Mulaku, Haddumati, Suvadiva, Kolumadulu and
North Male Atolls, in 25—42 fathoms.

28. Lambrus (Aulacolambrus) sculptus A. M.-Edw., 1872. Alcock, 1. p. 272 (Pl. XLVII.
fig. 6).
Dredged in South Nilandu, Felidu, Mulaku, and Suvadiva Atolls, in 20—43 fathoms.

29. Lambrus (Parthenolambrus) calappoides Ad. and Wh., 1847. Alcock, I. p. 275,
Dredged in South Nilandu and Suvadiva in 36 and 35 fathoms respectively.

XI. ON THE CLASSIFICATION AND GENEALOGY OF THE
REPTANT DECAPODS.

Besides the brachyurous Crabs, Boas’ Reptantia’ comprised sundry groups which are not
brachyurous, that is have a pair of biramous limbs on the sixth segment of the abdomen,
which bears traces of adaptation to other purposes than reproduction, such as swimming
or holding on a shell, and have also no fusion between the rostrum or front and the
epistome and no angle on the endopodite of the first maxilliped. The following are the
names of these groups: Eryonidea, Scyllaridea (= Loricata), Nephropsidea (= Homaridea and
Astacidea), Thalassinidea, Paguridea, Galatheidea, and Hippidea, the last three being together
known as Anomala by Boas and most other writers.

Our present object is to arrange these divisions according to their relationships, and to
do this we must first of all form an idea of the ancestor from which they may be supposed
to be descended. More space would be needed than can be spared here to set forth in
detail the process of comparison of the various types with one another and with the prawns
of the Stenopidea and Penaeidea, by which this result may be reached, and indeed this
might after all be hardly worth while, for the following statement will, I think, commend
itself to those who have studied the subject as on the whole probable.

The forebears of the reptant Decapods, if they could be examined, would probably show
the following characters :

(1) The rostrum of a good size, flat but keeled, and narrowly triangular.

(2) The carapace subcylindrical, free from the epistome both at the sides and in the middle,
overlapped behind by a process of the first abdominal segment on each side, and marked by the
following grooves: on the back two running transversely (the first, line e of Boas, is Bouvier’s

1 See above, Art. 1v. vol. 1. p. 424,

MARINE CRUSTACEANS. 691

cervical groove, and the second, line ¢ of Boas, is Bouvier’s branchial groove, ordinarily known as
the cervical groove; it would be well if these were known as the first and second cervical grooves);
at the sides three transverse grooves slanting forwards, the first two being prolongations of the
two cervical grooves and the third being line a of Boas; and two irregular longitudinal grooves, the
upper, line d of Boas, connecting the first cervical groove with the fore edge below the antenna,
and the second, line 6, 6’ of Boas, starting from the lower end of a@ and joining the lower ends

of the two cervical grooves with the fore edge some way below the line d’ (see Fig. 125).

(3) The abdomen long, straight, tapering somewhat from the second segment backwards,

moderately broad, not greatly compressed but with pleura bent
downwards and terga well arched. Both terga and pleura
stout and overlapping. The telson pointed.

(4) The eyes well developed and pigmented. The eye-
stalks of moderate length, made up of two cylindrical joints,

of which the second is longer than the first.

(5) The first antenna with a straight stalk of three
short, subequal subcylindrical joints, and two subequal flagella,
which are longer than the stalk, but shorter than the flagellum
of the second antenna.

(6) The second antenna with the basal joint free, a scale
which is broad but not leaf-like and ends in a sharp point
that represents the tooth on the outer angle in the prawns,
and a flagellum which is slender and about as long as the
body.

(7) ‘The mandible with a broad cutting edge, a small
stump to represent the molar process, and a stout, curved,
three-jointed palp.

(8) The first maxilla with the plate which represents
the basipodite broader than that which represents the coxo-
podite, and the endopodite of a good size, and divided into
two joints, the second of which is directed outwards.

(9) The second maxilla with the coxopoditic and _basi-
poditie plates both cleft, so that four lobes result, these lobes
all narrow, and the scaphognathite ending behind in an angle,
which is rather less than a right angle.

(10) The first maxilliped with a many-jointed flagellum
on the exopodite, a two-jointed endopodite which is not very
broad and an epipodite. The basipoditic plate deep, but not
broad.

Fic. 125. Cephalothoracic carapace of Callia-
nassa novaebritanniae from the left side,
1. first cervical groove—the line e of Boas
and ‘‘cervical groove” of Bouyier, 2. second
cervical groove—the line ¢ of Boas and
Bouvier’s ‘branchial groove,” ordinarily
known as the ‘‘ cervical groove,” 1’ and 2’.
continuations of 1 and 2 at the sides of
the carapace, 3. upper longitudinal line—
the linea thalassinica and perhaps also the
linea homolica—the front part of which is the
line d of Boas, 4. lower longitudinal line—
the linea anomurica and perhaps also the
linea dromiidica—the front part of which is
the line b of Boas, 5. a soft area at the side
of the carapace, 6, 6’. hard plates in this
area, 7. rostrum, 8. thickened hinder edge of
the carapace. The second cervical groove is
here represented only by a line or crack in
the carapace. It is, however, in the same
place as the cervical groove of other Callia-
nassas, and there is a good deal of evidence
that a groove may be represented by a
erack, The linea thalassinica, for instance,
is sometimes a groove in part of its length.

(11) The second mawilliped with the exopodite longer than the endopodite, carrying a jointed
flagellum, and the endopodite slender, with the last joint at the end.

1 A backward prolongation of the upper of these lines (d),
known as the linea thalassinica, is found in many Thalas-
sinidea, and a similar prolongation of the lower (b), known
as the linea anomurica, in the Anomala, The linea homolica
and linea dromiidica correspond in position with the first

and second of these respectively, but as they are not con-
tinuous with d and b it is doubtful how far they can be looked
upon as homologous.
loosely applied to the longitudinal suture of the Homolidae
(as on p. 575 above) is wrong.

In any case the term linea anomurica

692 L. A. BORRADAILE.

(12) The third masilliped with the exopodite shorter than the endopodite, carrying a straight,
jointed flagellum directed forwards, and the endopodite narrow, with separate ischiopodite and
meropodite.

(13) All the Jegs seven-jointed. The first three pairs chelate with normal chelae, the first
the greatest, equal, the fourth pair simple, the fifth pair simple (?) and having its last two joints
slightly twisted so that, when the basal joints are pressed against the sides of the body, while
the last joint of the fourth pair points downwards and backwards, that of the fifth pair points
forwards and inwards’.

(14) The gills many, including mastigobranchs and podobranchs on all the thoracic limbs but
the last, arthrobranchs on all but the first and last, and pleurobranchs on the last four.

(15) The thoracic segments which bear the legs with distinct sterna, which are not very broad,
but grow broader from before backwards, and are all fused except the last”.

(16) The first pair of abdominal limbs unbranched. Those of the second to fifth segments
with two fairly broad branches, the inner of which bears an appendix interna (Fig. 126c). The last
pair about as long as the telson, with broadly oval branches, across the outer of which is a suture.

The internal anatomy of the various groups is not yet well enough known to allow of general
statements being made about it.

The animal was hatched in the Zoea stage, with a segmented abdomen but no limbs behind
the third maxilliped, and passed through a Mysis stage with exopodites on all the legs.

The descendants of the crustacean which we have thus reconstructed fall into two
sets, one comprising the Eryonidea, Scyllaridea, and Nephropsidea, in which the abdomen
is strong and well-armoured, stretched out unprotected, and used as a swimming organ by
means of its tail-fin, and the other containing the rest of the groups, in which it has for
some reason become a burden and a source of danger to be protected and kept from
exposure, even though it be still shaped and used for swimming. The first of these sets
is on a lower and more primitive grade of organisation than the second. This is shown
(1) by the abdomen, with its stout armour, overlapping terga and pleura, strong processes
to clip the carapace on the first segment, broad tail-fin, and, in the Eryonidea and Scyllaridea,
appendices internae. To this form of abdomen, which is always carried at length, I shall
restrict the term “macrurous.” (2) by the legs, which in the Eryonidea and Nephropsidea
are chelate in the first three pairs and have seven joints (except for the first pair of the
Nephropsidea) whereas in all other Reptantia they have only six, owing to the fusion of
the basipodite and ischiopodite, (3) by the large number of the gills, (4) by the slender
third maxillipeds with their long flagella directed forwards, (5) by the broad antennal scale of
the Eryonidea and Nephropsidea. Of course some of the foregoing characters are found in
primitive members of other groups, but they stamp this set of groups as a whole. Of the
three macrurous groups, the Eryonidea and Scyllaridea are more nearly allied together than
either of them is to the Nephropsidea. They have both lost, or much reduced, their rostrum
(except Palinurellus), reduced the inner lobes of the second maxilla, and fused their carapace
at the sides with the epistome, but they have kept the appendices internae. Their body

1 This character, though it is most prominent in the but also in the Thalassinidea, Potamobiidae and Parastacidae
Anomala and Thalassinidea and the lower Crabs, is seen to and in the prawn Stenopus. In Penaeus the anterior sterna
some extent even in the more primitive groups, as the are free, but the last two are joined by secondary thickenings
Palinuridae and Potamobiidae. in the membrane between them.

2 A primitive feature, found not only in Boas’ Anomala

MARINE CRUSTACEANS. 693

shows a tendency to be flattened. The Nephropsidea differ from them in all these respects.
Between the Eryonidea and the Scyllaridea the most striking difference lies in the fact
that in the former all the legs are chelate and in the latter none are so. The Eryonidea
are the more primitive in keeping the antennal scale, the joint between the ischiopodite
and meropodite, and the chelae on the first three legs.

Fie. 126. Limbs of Avius acanthus; a, second maxilla, b. first maxilliped, c. abdominal limb of the third pair.
1. gnathobase or first lobe of coxopodite, 2. endopodite, 3. appendix interna, 4. lobes of basipodite.

In considering the second set of groups, we have first to settle whether to look upon them
as of common descent and treat them as a whole, or whether we must find a separate point
of origin for each of them from among the Macrura. Now it may be allowed at the outset
that there is but one character that every member of them shares with the rest. In the
macrurous groups the first three pairs of legs are all alike, adapted either to seizing or to
crawling—chelate, that is, or simple. But in the modified groups we are now dealing with
the third pair is never adapted to the same end as the first. In the overwhelming majority
the first pair are chelate or subchelate’ and the third used for walking, and in the Hippidae,
where the first are simple and straight, the third are hooked and flattened for burrowing or
swimming. Nor is this point an unimportant one, for the likeness of the first three legs
is a heritage from the penaeid prawns, and as such is of considerable value. Moreover it
is possible to find a number of characters which, while they are not seen in every member
of the non-macrurous groups, are yet common in each of them, and do not occur elsewhere.
These are: (1) a large gnathobase, or inner coxopoditic lobe, on the second maxilla (Fig. 126 a),
(2) the shape of the first maxilhped, which has a broad but shallow basipoditic lobe, whose
inner margin is in the same line with that of the coxopodite*®, and a broad basal half to
the exopodite (Fig. 126 6), (8) the flagella of the maxillipeds turned inwards and curved
forwards at their ends, (4) reduction in various ways of the abdomen, (5) the presence of
backward prolongations of the longitudinal lines of the carapace*. Since, however, numbers
(8) to (5) are less strongly marked in the more primitive forms of some of the groups, it
is doubtful whether much stress can be laid upon them.

Taking it for granted, then, that the non-macrurous Decapods had a common, if remote,
ancestor, it remains to be seen where that ancestor must be placed with regard to the

1 Gebicula seems at first sight to be an exception to this, ‘‘ thumb” of the allied Upogebia.
but there is on the propodite a strong tooth near the end * Found also in at least one species of Nephropsis.
which seems almost certain to be used in the same way as the 3 See above, footnote to p. 691.

G. Il. 89

694 L. A. BORRADAILE.

macrurous groups. For this purpose we must add to the list above certain other characters
which, either from their own primitive character or because they are found in many of the
primitive members of the groups we are trying to place, may be attributed to the forebears
of these groups: (6) a large gill-formula, including all the gills of the ancestral reptant’,
(7) a small thorn-like antennal scale?, (8) the last pair of legs not only on a free sternum®
as in the Potamobiidae and Parastacidae, and slightly twisted at the end as in most
macrurous Reptantia, but distinctly smaller than the fourth pair, removed from them, and
carried more dorsally, (9) a rather short and broad rostrum, (10) appendices internae on
abdominal limbs’, (11) a transverse suture on the telson‘.

Numbers (6) to (9) of these remove the crustaceans in which they are found from the
neighbourhood of the Eryonidea and Scyllaridea, since they are primitive features that the
latter have lost. Numbers (6) and (10) remove them for the same reason from the
Nephropsidea, to which, nevertheless, they are more akin than to the other macrurous groups.
The jaws and the last thoracic sternum, but not the structure of the gills or the grooves
on the carapace, are more like those of the Potamobiidae than those of the Nephropsidae.

Assuming, as above, a common descent for the Crabs, Anomala and Thalassinidea, we
must at the same time admit that they very early divided into two widely separated
branches, one containing the Crabs and the other the remaining groups. The characters by
which this separation is shown are the following: (1) The great reduction of the abdomen
in the Crabs. This is entirely independent of similar developments in the higher Anomala,
for the more primitive of the Paguridea and Thalassinidea, though they share the tendency,
common to the whole assemblage of families under discussion, to shelter and protect their
abdomen, have as yet been hardly more affected by it than, say, the burrowing genera of
Nephropsidea, which are quite macrurous, while the primitive Galatheidea are little better
in this respect. Moreover, in the Crabs, the reduction has gone so far that the abdomen,
having lost its sixth pair of limbs (except for doubtful, unbranched vestiges in some Dromiidea),
is now fitted for, and shows traces of. no other function than those connected with repro-
duction, while in the Anomala and Thalassinidea, though some of its macrurous features are
always reduced, it has kept the sixth pair of limbs and nearly always uses them either for
swimming or for holding on a shell. The Lithodinea alone form an exception to this state-
ment, but in them the asymmetry of the abdomen clearly recalls its former use, as in the
Hermit crabs, to hold on a shell. These two types of abdomen—of which one does, while
the other does not, show traces of adaptation to some other function than that of reproduction
—I propose to call the “brachyurous” and “‘anomurous” respectively. (2) The carapace, which
in the Anomala and Thalassinidea remains free, is in the Crabs fused with the epistome both
at the sides and (except in Homolodromia) in the middle, under the front. (3) The antennal
scale, which in many Anomala and Thalassinidea remains and is moveable, is never found in
that condition in the Crabs. (4) The endopodite of the first maxilliped is broad and has
nearly always an outer angle in the Crabs (see above, p. 425, fig. 110), but has not this shape
in the Anomala and Thalassinidea, though some Hippidea approach it. (5) There are never
appendices internae on the abdominal limbs of the Crabs, whereas these structures are present
in most Thalassinidea and some Anomala. (6) The third pair of maxillipeds of the Crabs are

1 See above, p. 692. This is shown by Jaxea, Homolo- is fused to the stalk.
dromia, ete. 3 Not found in the Crabs.
2 Among the Crabs found only in Homolodromia, where it 4 Found in certain Galatheidea and Paguridea.

MARINE CRUSTACEANS. 695

usually broad, forming a cover to the mouth, whereas those of the Thalassinidea and Anomala
are only exceptionally so. These characters, however, are in themselves hardly enough to
negative the supposition that the Crabs have originated from one of the other non-macrurous
groups. What does make it impossible to derive them from the Anomala is the presence
in the primitive crab Homolodromia of podobranchs on some of the legs, while an origin
from the Thalassinidea is equally untenable on account of the less primitive condition of the
grooves of the carapace and the reduction of the endopodite and flagellum of the first
maxilliped in the latter group.

We are now left with the Thalassinidea and the three divisions of the Anomala. still
unaccounted for. Among these there can be no question that the Thalassinidea are, as a
whole, the most primitive. Their straight symmetrical abdomen, with well-developed biramous
limbs on all the segments except the first, and the sixth pair broad and flat in all but
Thalassina, the presence in most of appendices internae, and in some of good pleura, the
simple legs of the fourth pair and sometimes also of the fifth, the rostrum usually well-
developed, the moveable antennal scale seldom wholly lost, and the large gill-formulae of
many genera, extending from the second maxilliped to the fourth leg, with mastigobranchs,
podobranchs, arthrobranchs and sometimes pleurobranchs

all these point to the same con-
clusion. Nor can there be any doubt that the Paguridea and the Thalassinidea are closely
akin and have branched off from the same not very remote ancestor. The only differences
between the Axidae and the primitive Pagurids with symmetrical abdomen such as Pylocheles
are the presence of a pleurobranch on the last leg, a suture across the telson in some species
(which is never found in Thalassinidea), better developed eyes!, the loss of epipodites on the
legs, a reduced rostrum, subchelate legs of the fourth pair, and the branches of the last pair
of abdominal limbs narrow and not adapted for swimming. The first three of these features,
being primitive, show that the Paguridea are not to be derived from any of the existing
Thalassinidea and the remainder make it equally impossible to derive the latter group from
the former. The two meet at a point below our present horizon.

The Galatheidea join the stem from which the last two groups arise before its
bifurcation, that is, they are less closely akin to either of these than the latter are to one
another. The fact that they may have epipodites on some of the legs, and the more primitive
shape of the rostrum and sixth pair of abdominal appendages make it impossible to place
their ancestor within the present limits of the Paguridea. The suture on the telson and
the pleurobranch on the last leg remove it also from the Thalassinidea. And the general
shape of the body, depressed, with broad flat abdomen carrying long pleura and bent under
the thorax, is so different from the compressed body, with straight abdomen, of the primitive
Paguridea and Thalassinidea that there can be little doubt that the ancestral Galatheid left
the non-macrurous stock, after the Crabs indeed, but before it gave rise to the Thalassinids
or Hermit-crabs. As for the linea anomurica, this is found not only in the Galatheidea and
Paguridea but also very distinctly in Callianassa novaebritanniae, and it is curious to notice
that it is wanting in the primitive Pagurids such as Pylocheles, and most Thalassinids as in
Azius, and that in the Galatheid Aeglea, which is also primitive in many respects, the hinder
part is again wanting. Thus it would seem as though this line appeared only in the higher
and more typical forms of each group. The Hippidea present a very difficult problem. On

1 Such species as Awiopsis clypeatus, however, have the eyes well developed, though not so large as those of the
Paguridea.

89—2

696 L. A. BORRADAILE.

the whole, however, their general shape of body, with a bent abdomen carrying good pleura,
subeylindrical or depressed cephalothorax, and fourth pair of legs like the third rather than
the fifth, shows a nearer kinship to the Galatheidea than to any of the other groups, and
in the absence of any evidence to the contrary this judgment must stand. As for their
likeness to the Raninidae among the Crabs, an admission that this indicated relationship would
lead to great difficulties, as for instance that the Raninidae must either be removed from
the closely related Oxystomes or the latter be supposed to be derived by modification from
the highly specialised Hippidea—in itself a very unlikely speculation.

Each of the above groups has its peculiar habits and habitat. The Galatheidea hide
under stones or dwell in weed. The Thalassinidea generally make burrows, but the more
primitive genera among them show a tendency to shelter in weed or sponges. The Paguridea
(except Lithodinea) place their abdomen in the hollow of some foreign body, which is usually
a gastropod shell, but in the primitive genera may be a stone or a sponge. The Hippidea
bury themselves in the sand. Of these habits, that of the Galatheidea is the least specialised
and could easily have given rise to the others.

We are now in a position to sum up in the form of a tree the results reached in this
and former articles (Pl. XLVIII.). When this is done two facts of importance appear. First
that the proper place of the Thalassinidea is in the midst of the anomurous groups, and
that they must no longer be classed with the Macrura, and secondly that, when this change
is made, the Anomura, like the Brachyura, become a true, monophyletic group. Thus the
reptant decapods fall into three divisions, Brachyura, Anomura’ and Macrura. To the subject
of the taxonomic value of these groups and their relation to the natant families I hope to
return later on. For the present they may be called suborders’.

The following keys give more precisely the classification of the groups discussed above :

Key to the reptant suborders of the Decapoda.

I. 3rd pair of legs like the first, either chelate or simple and subcylindrical. Abdomen macrurous
(straight, symmetrical, well armoured, with good pleura and strong, broad tail-fin, a lobe on
the first segment clipping the carapace). Gnathobase of 2nd maxilla narrow. Basipoditic lobe
of lst maxilliped usually deep. Exopodites of maxillipeds with lash directed forwards. Gills
numerous. [Last thoracic segment with legs not differing greatly from the rest and sternum
rarely free. | Macrura.

II. 3rd pair of legs unlike the first’, never chelate. Abdomen rarely macrurous. Gnathobase of
2nd maxilla typically broad. Basipoditic lobe of Ist maxilliped broad but shallow, its inner
edge in a line with that of the coxopodite. Exopodites of maxillipeds with lash, when present,
nearly always bent inwards. Gills usually few. Last thoracic segment with limbs often differing
greatly from the rest and sternum free or not.

1. Carapace not fused with epistome. Last thoracic sternum free, its legs differing always clearly
in size and position and nearly always in shape from the third pair. Abdomen anomurous
(reduced in some of its features but showing clear traces of some other function than that
of reproduction and almost always carrying biramous limbs on the 6th segment) or, rarely,

1 Unlike Boas’ term ‘‘ Anomala,” the name ‘‘Anomura” which it has not had before.
has had yery different meanings in the pages of different 2 The Brachyura is ranked lower on p. 427.
writers. This fact allows it to be used here with a denotation 3 See footnote to p. 693.

MARINE CRUSTACEANS. 697

macrurous. A moveable antennal scale often present. Third pair of maxillipeds usually
narrow. Anomura.

2. Carapace fused with epistome at the sides and nearly always in the middle. Last thoracic
sternum fused with the rest, its legs often like the others. Abdomen brachyurous (small,
straight, symmetrical, bent under the thorax, showing no traces of any other function than
that of reproduction, and without biramous limbs on the 6th segment). Never a moveable
antennal scale. 3rd pair of maxillipeds usually narrow. Brachyura'.

Key to the tribes of the reptant Macrura.

I. Carapace fused at the sides to the epistome. Rostrum small or wanting (except Palinwrellus).
Inner lobes of 2nd maxillae and Ist maxillipeds reduced. An appendix interna on some of
the abdominal limbs, at least in the female, but the exopodite of the last pair without sharp
suture. Body often depressed.

1. Carapace gripped by first abdominal segment alone. First joint of antenna not fused with
epistome ; a scale present on this limb. All the legs, except sometimes the last pair, chelate;
the first longer than the rest. Unbranched limbs on the first abdominal segment. Tail-fin
not softer behind than before, without sutures. Telson pointed. Eryonidea.

1)

Carapace gripped between a lobe on the lst abdominal segment and a knob on the side
of the last thoracic segment. First joint of antenna fused with epistome; no scale on this
limb. None of the legs much longer than the rest or, except sometimes the first pair,
chelate. No limbs on the Ist abdominal segment. Tail-fin divided by indistinct sutures
into a soft hinder half and a harder front half. Telson roughly squared behind. Scyllaridea’.

II. Carapace free from the epistome. Rostrum of good size. Inner lobes of 2nd maxillae and
Ist maxillipeds not reduced. No appendix interna, but the exopodite of the last abdominal
limb divided by a suture. Body subcylindrical. [Carapace gripped by first abdominal segment
only. Ist joint of antenna free. A scale present. First three legs chelate, first pair the
longest. Telson firm, squared, often sutured. ] Nephropsidea.

Key to the tribes of the Anomura.

I. End-joints in 2nd to 4th legs curved and flattened. First pair styliform or subchelate. [Tail-fin
not adapted for swimming. Abdomen bent under thorax. Rostrum small or wanting. 3rd pair
of maxillipeds carry no mastigobranchs. | Hippidea’.

II. End-joints in 2nd to 4th legs not curved and flattened. First pair not styliform, rarely

subchelate.

1. 6th abdominal limb adapted for swimming (except in TZhalassina where it is styliform.
Pleura usually well developed. Abdomen symmetrical.

a. Body depressed. A pleurobranch to the last leg. Often a transverse suture on the
telson. Abdomen more or less bent under the thorax, Galatheidea.

6. Body compressed. No pleurobranch to the last leg. No transverse suture on the telson.
Abdomen straight. Thalassinidea*.

1 For a classification of the Crabs, with keys, see above, articles which deal with them.
vol. 1. pp. 426 ff. 3 For a classification of this group, with keys, see Ann.
2 Keys to the families of these groups will be givenin the Mag. N. H. (7), xu.

698 L. A. BORRADAILE.

2. 6th abdominal limb, when present, has the branches neither broad nor styliform but adapted
for holding the body into hollow objects. Pleura very rare. Abdomen nearly always
asymmetrical and either soft and twisted, or bent under the thorax. Paquridea.

Key to the subtribes of the Paguridea.

I. Abdomen straight or twisted. Carapace firm and more or less compressed in the fore part,
soft in the hinder part. Fourth pair of legs unlike the third. Rostrum almost or quite
wanting. Pagurinea.

II. Abdomen bent under the thorax.- Body crab-like. Carapace firm all over. Fourth pair of
legs like the third. Rostrum spiniform. Lithodinea.

EXPLANATION OF PLATE XLVII.

Fic. 1. Huenia proteus, male of triangular shape; a. whole animal, 5. part of the fine graving
of the back, greatly magnified.

Fie. 2. Huenia proteus, female.

Fic. 3. Naxioides spinigera, male; a. whole animal, }. outside of hand, ¢. hooked hairs of two
different shapes.

Fic. 4. Halimus espinosus, male; a. whole animal, 6, outside of hand, c. end-joint of a walking leg,
d. hooked hairs.

Fie. 5. Lambrus (Rhinolambrus) turriger; a. whole animal, 6. part of under side to show grooves
for breathing stream. 1. endopodite of 3rd maxilliped, 2. exopodite of the same, 3. base of
epipodite of the same, covering opening to gill-chamber, 4. ridge on under side of body (not
outer edge), 5. outer end of inner groove, 6. outer end of outer groove, 7. inner groove, 8. fore
edge of mouth,

Fic. 6. Lambrus (Aulacolambrus) sculptus; part of inner groove corresponding to (7) in fig. 5,
showing the covering of hairs.

Fauna and Geography, Maldives and Laccadives Plate XLVII

E. W. del.

BORRADAILE——OXYRHYNCHA,

Plate XLVIII

Fauna and Geography, Maldives and Laccadives

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Now ply ayy ‘apoll snopoumowp ay? UWL alM Mol Pwov—as ay? WL ASOYT, “SQN. ayy sD uounsruvhio fo appub aus ayy payavat 7)0 aany Now qgsuf ayy ua sdnowb aT,

‘yaOdvVOad INV1IddaY S3HL SO 351 TWOISOIVANAD

(VaAMOVN)

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anpisdo..yda ay

INPYOYIO SOUND YT,
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appriqout DOI

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(VLVWOISAXQ) (VIOVINOU(] )(VHLVNOAHOVU )
Ya

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” 76 yea ADPULO.UpO]OULo FT

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Now ready, Parts I—lV forming Volume T.

The Fauna and Geography
of the
Maldive and Laccadive Archipelagoes

Being the Account of the Work carried on and
of the Collections made by an Expedition
during the years 1899 and 1900

Edited by

J. STANLEY GARDINER, M.A.

Fellow of Gonville and Caius College and late Balfour Student
of the University of Cambridge.

VOLUME I.
With Plates I—XXV and Text-Illustrations 1—119

CAMBRIDGE :
at the University Press.

Lonpon: C. J. Ciay anp Sons,
Cambridge University Press Warehouse,
Ave Maria Lane.

GLascow: 50, Wellington Street.

1903

Volume I. of the above work is now complete and contains the
following Reports :—

il

10.

hile

r

13.

14.

15.

16.

Introduction: Narrative and Route of the Expedition. With
Text-Figs. 1 and 2

By J. Santry GARDINER, M.A.

The Maldive and Laccadive Groups, with Notes on other Coral
Formations in the Indian Ocean. Chapters I—IV. With
Plates I and II and Text-Figs. 3—11

By J. Stanney GARDINER, M.A.

Hymenoptera
By P. CAMERON.
Land Crustaceans. With Plate III and Text-Figs. 12—23
By L. A. Borrapaite, M.A., Lecturer in Natural Sciences of Selwyn College,

Cambridge.
Nemerteans. With Plates IV and V
By R. C. Punnett, B.A.
Amphibia and Reptilia
By F. F. Larpuaw, B.A.
Lepidoptera ;
By Ep. Meyrick, B.A., F.Z.S.
Echiuroidea. With Plate VI
By A. E. Surpiey, M.A.

Sipunculoidea, with an Account of a new Genus Lithacrosiphon.
With Plate VII
By A. E. Saipiry, M.A.

Land and Freshwater Mollusca. With Text-Fig. 24

By EpeGar A. Smiru.

The Maldive and Laccadive Groups, with Notes on other Coral
Formations in the Indian Ocean (continued). Chapters V—VII.
With Plates VIII—XII and Text-Figs. 25—33

By J. StanLEy GARDINER, M.A.

On the Pigments of certain Corals, with a Note on the Pigment
of an Asteroid. With Text-Fig. 34
By C. A. MacMunn, M.A., M.D.

Marine Crustaceans. I. On Varieties. II. Portunidae. With
Text-Figs. 35—38
By L. A. Borrapaiie, M.A.

Chaetognatha, with a Note on the Variation and Distribution
of the Group. With Plate XIII and Text-Figs. 39 and 40

By Lronarp Doncaster, B.A.

Dragon-Flies
By F. F. Larpnaw, B.A.
The Actinogonidiate Echinoderms of the Maldive and Laccadive

Islands : :
By Prof. F. Jerrrey Beni, M.A.

PAGE

12

51

64

101

119

123

127

131

141

146

184

209

219

223

ie

18.

il).

22.

23.

24.

25.

26.

27.

28.

CONTENTS.

Orthoptera : :
By Matcoutm Burr, F.Z.S., ete.

Marine Crustaceans. III. The Xanthidae and some other Crabs.
With Text-Figs. 41—60
By L. A. Borrapatie, M.A.

On the Fishes from the Maldive Islands. I. Dredged. II. Fresh-
water .
By C. Tare Reean, B.A.

The Marine Turbellaria, with an Account of the Anatomy of
some of the Species. With Plates XIV and XV and Text-
Figs. 61—73 : ‘ ‘ : 5 : é

By F. F. Larnaw, B.A.

The Maldive and Laccadive Groups, with Notes on other Coral

DRAPE HHH He MOTE E TE SHEE SHES SEOSHSESHESHEHHEEEHEOTEHSES eereseee

Formations ‘in the Indian’ ‘Ocean’ (continied). “Appendix ‘A.
With Plates XVI and XVII and Text-Figs. 74 and 75.

By J. Sranney GarpINER, M.A.

Cephalochorda.

Systematic and Anatomical Account. With Plate XVIII and
Text-Figs. 76—89 .
By C. Forster Cooper, B.A.

II. Note on Meristic Variation in the Group

By R. C. Punnett, M.A.

Aves : : ‘ : P ;
By H: Gapow, M.A., F.R.S., and J. Sranney Garpiner, M.A.

The Earthworms of the Maldive and Laccadive Islands
By Frank E. Bepparp, F-.R.S.

The Maldive and Laccadive Groups, with Notes on other Coral
Formations in the Indian Ocean (concluded). Appendices B

and C. With Plates XIX—XXI and Text-Figs. 90—109

By J. Srantey GarpIner, M.A.

Marine Crustaceans. IV. Some Remarks on the Classification
of the Crabs. V. The Crabs of the Catometope Families.
VI. Oxystomata. VII. The Barnactes. With Plate XXII
and Text-Figs. 110—119 . 2 : ; 3 ;

By L. A. Borrapaine, M.A.

Marine Crustaceans. VIII. Stomatopoda, with an Account of
the Varieties of Gonodactylus chiragra. With Plate XXIII .
By W. F. Lancuester, M.A.

The Lithothamnia of the Maldives and Laccadives. With Plates
XXIV and XXV
By M. Fostir.

PAGE

234

237

272

282

313

347

368

374

376

424

444

460

It was supposed that the whole work would be completed
in eight parts, forming two volumes, but it is now anticipated
that one or more supplementary parts may have to be issued ;
each part will contain about 120 pages of text, demy quarto,
and will be fully illustrated by lithographic plates and _ text
illustrations prepared by Mr Epwin WiLson of Cambridge.

The edition is limited to 475 copies and the subscription
for the whole work including any supplementary parts that
may be issued is for the present £4 4s. net. The separate parts
are published at 15s. net.

FORM “OF SUBSCRIPTION:

Messrs*C.) J i@ray cz Sons;
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Ave Maria Lane, London.

Please send me 22. = Op Of

The Fauna and Geography of
the Maldive and Laccadive Archipelagoes

price £4 4s. per copy (¢ucluding any supplementary parts). The

work to be forwarded in separate parts on publication.

WN OMEC se 4 got ge ars Pk a ee hee

ACAIESS he ON es