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PROCEEDINGS

of the

Biological Society of
Washington

VOLUME 95
1982

Vol. 95(1) published 13 April 1982 Vol. 95(2) published 11 August 1982
Vol. 95(3) published 5 October 1982 Vol. 95(4) published 20 December 1982

WASHINGTON
PRINTED FOR THE SOCIETY

EDITOR

BRIAN KENSLEY

ASSOCIATE EDITORS
Classical Languages Invertebrates
GEORGE C. STEYSKAL THOMAS E. BOWMAN
Plants Vertebrates
DAVID B. LELLINGER RICHARD BANKS
Insects

ROBERT D. GORDON

All correspondence should be addressed to the
Biological Society of Washington, Smithsonian Institution
Washington, D.C. 20560

ALLEN PREss INC.
LAWRENCE, KANSAS 66044

OFFICERS AND COUNCIL
of the
BIOLOGICAL SOCIETY OF WASHINGTON
FOR 1981-1982

OFFICERS
President
RAYMOND B. MANNING

Vice President
PAUL J. SPANGLER

Secretary
MICHAEL A. BOGAN

Treasurer
LESLIE W. KNAPP

COUNCIL
Elected Members
FREDERICK M. BAYER KRISTIAN FAUCHALD
ISABEL C. CANET DAVID L. PAWSON

AUSTIN B. WILLIAMS

TABLE OF CONTENTS

Volume 95

Adis, Joachim, and Richard C. Froeschner. Notes on distribution of some Latin Amer-
ican cotton-stainers (Dysdercus: Pyrrhocoridae: Hemiptera) and remarks on the biol-
ogy of Dysdercustunbahnt- Schmidt. .2 ee ee ree ep ase pee e

Adkison, Daniel L. Description of Dactylokepon sulcipes n. sp. (Crustacea: Isopoda:
Bopyridae)randsnotesiOmel) Cari Ge See cee era ee ee ae eee ee

Adkison, Daniel L., Richard W. Heard, and Guy T. Clark. Description of the male and
notes on the female Argeiopsis inhacae (Crustacea: Isopoda: Bopyridae) ______________

Baker, H.R. A note on the genitalia of Potamothrix hammoniensis (Oligochaeta: Tubi-
TGUAAE) i se a ee ec a

Barnard, J. Laurens, and Margaret M. Drummond. Redescription of Exoediceros fossor
(Stimpson, 1856) an Australian marine fossorial amphipod, the type-genus of the new
family EP xoedicerotidac \.. 2-5 2 Be ee

Barnard, J. Laurens, and Gordon S. Karaman. Classificatory revisions in gammaridean
Amphipoda (Crustacea) 3 Pamt.2.¢ 202° 2k NS ae ie Mine ee eee dr se ee

Bayer, Frederick M. Some new and old species of the primnoid genus Callogorgia
Gray, with a revalidation of the related genus Fanellia Gray (Coelenterata: An-
tHOZOB) ee a wee eT Ne en Oe

Bowman, Thomas E., and Richard Franz. Anopsilana crenata, a new troglobitic ciro-
lanid isopod from Grand Cayman Island, Caribbean Sea ______________________________-_-

Brooks, Daniel R., and Janine N. Caira. Atrophecaecum lobacetabulare, n. sp. (Di-
genea: Cryptogonimidae: Acanthostominae) with discussion of the generic status of
Paracanthostomum Fischthal and Kuntz, 1965, and Ateuchocephala Coil and Kuntz,
| | en eee Ome eee Me Cen ee ee

Brown, Walter C., and Angel C. Alcala. A new cave Platymantis (Amphibia: Ranidae)
from, the Philippine Islands)... 0 a eo

Bruce, Niel L., and Thomas E. Bowman. The status of Cirolana parva Hansen, 1890
(Crustacea: Isopoda: Cirolanidae) with notes on its distribution __________________________

Chernoff, Barry, and Robert Rush Miller. Mexican freshwater silversides (Pisces: Ath-
erinidae) of the genus Archomenidia, with the description of a new species ____________

Child, C. Allan. Pycnogonida of the western Pacific islands I. The Marshall
| E10 | ea ieee ee ee coe eR OAD Ay ge om cce te) Sat ee

Child, C. Allan, and Koichiro Nakamura. A gynandromorph of the Japanese pycnog-
onid=Anoplodaciylus ge'stien's | (@rctriverramn)) seme eee eae ea ee een

Collette, Bruce B. South American freshwater needlefishes of the genus Potamorrha-
phis (Beloniformes: Belomidac)y eae: eee ee ee ee See ee

Cressey, Roger F. A new genus of bomolochid copepods from Indo-West Pacific nem-
UPterid fishes = Sebo os os EN ir ce an fk, Sole sks lee! lee

Crews, Celinda R., and Artie L. Metcalf. A new species of oreohelicid land snail from
the: Sam “Agus Gime plains INS wi VIS xc sae a i ne

Cutler, Edward B., and Norma J. Cutler. A revision of the genus Siphonosoma
(Stpuncula): 225 20.24 pack te ew SN pen ger angel een lr ie

Dawson, C. E. Review of the genus Micrognathus Duncker (Pisces: Syngnathidae),
with descriptionrof iM. satanis me Spee aasake nee ae setae ee nse an eee te) eee

Dawson, C. E., and C. J. M. Glover. HAypselognathus horridus, a new species of
pipefish (Syngnathidae) from South Australia _____.-________-_------ Sessa

Desbruyéres, Daniel, and Lucien Laubier. Paralvinella grasslei, new genus, new species
of Alvinellinae (Polychaeta: Ampharetidae) from the Galapagos Rift geothermal
VMS) ce ue Ee rai i a RT Soe ent ae oie gas ail

Downey, Maureen E. Evoplosoma virgo, a new goniasterid starfish (Echinodermata:
Asteroidea) from the:Gulf of Mexico. 2 2s eee

Ewing, R. Michael. A partial revision of the genus Notomastus (Polychaeta: Capitel-
lidae) with a description of a new species from the Gulf of Mexico ____-_________________

iv

371-376
702-708
334-337

563-566

610-620

167-187

116-160

522-529

223-231
386-391
325-333
428-439
270-28 1
292-296
714-747
495-504
256-264
748-762
657-687

403-407

484494
772-173

232-237

Fauchald, Kristian. Two new species of Onuphis (Onuphidae: Polychaeta) from Uru-
ON hy NE NS Ned Nin Sp ates NY ed Bere A an Pk Ne as Na AG ooh ed | Se jt ard
Fauchald, Kristian. Some species of Onuphis (Polychaeta: Onuphidae) from the Atlan-
(WE QUESBIN » wae creitich ME tae 2 halen iia Beek SE Teal, ie epee N Nc, bs Bigs ABNER lige se Se Ree 1 ae WE Reea aN er a
Fauchald, Kristian. A eunicid polychaete from a white smoker __________________-__________
Fauchald, Kristian. Description of Mooreonuphis jonesi, a new species of onuphid poly-
chaete from shallow water in Bermuda, with comments on variability and population
EC iy me ne EL eked ML EAN yl seweN Tee Ny I AEE 2 vera Enthe Usb net S
Flint, Oliver S., Jr., and Joaquin Bueno-Soria. Studies on Neotropical caddisflies, XXXII:
the immature stages of Macronema variipenne Flint & Bueno, with the division of
Macronema by the resurrection of Macrostemum (Trichoptera: Hydropsychi-
TEC) EL he apa) Su sl eve eae pe Nc babe Ll eee Boo dele
Formas, J. R., and Alberto Veloso. Taxonomy of Bufo venustus Philippi, 1899 (Anura:
Peomiodacmidae)mnom, central Chile... 2 22... 3.50 se
Formas, J. R., and M. Ines Vera. The status of two Chilean frogs of the genus Eup-
NepuemeAnunacmeeplodactylidac) .-.. 8.2. 22.2 ee ee a BL
Fredette, Thomas J. Evidence of ontogenetic setal changes in Heteromastus filiformis
duolehactan@apivellidae)irummr. Sa Soe meal aiel! yale 6 ok eb wl eat ol
Garcia, Renato G., and Raymond B. Manning. Four new species of stomatopod crus-
Tee SmnROMMecOe EOP pines 2.26 eee ee ee ee ees
Garrido, Orlando H., and Albert Schwartz. A new species of Sphaerodactylus (Reptilia:
SanasGekkonidae) from eastern Cuba 222) 2
George, Robert Y., and Noel A. Hinton. A new species of deep-sea isopod, Storthyn-
gura myriamae, from the Walvis Ridge off South Africa __________-____-_--_---
Gleye, LindaG. Two new species of leptomysinid mysids (Crustacea: Mysidacea) from
SOURS (CANINE: «SUIT RN sh oe eM eke tags BR et eemmeeane y cNeeS ORs bieaeeae aL seem ee
Green, Karen D. Uncispionidae, a new polychaete family (Annelida) ____________________
Harasewych, M. G. Pterynotus xenos, a new species of muricid from off northern
Teamraneancwionnscas Gastropoda), 22.2209 se2e8 8 ee ee
Harding, Keith A. Courtship display in a Bornean frog _____________________-______-----_-_-
Heaney, Lawrence R., and Gary S. Morgan. A new species of gymnure, Podogymnura
(Mammalia: Erinaceidae) from Dinagat Island, Philippines ____________-____--__-__--
Hershkovitz, Philip. Subspecies and geographic distribution of Black-mantle Tamarins
NOCnmUmsS Mericolis Spix (Primates: Callitrichidae) ___..._...-.-..2- 2-222. ee
Heyer, W. Ronald. Two new species of the frog genus Hylodes from Caparao, Minas
Gemiss braziy (Amphibia: Weptodactylidae) 2220..00 82.2). e ee ee
Heyer, W. Ronald, Charles H. Daugherty, and Linda R. Maxson. Systematic resolution
of the genera of the Crinia complex (Amphibia: Anura: Myobatrachidae) ______________
Hobbs, Horton H., Jr., and Daniel J. Peters. The entocytherid ostracod fauna of north-
Saar 21 hl es SUES elie bin). 9a he be cy) pe do =O ht hae od DY ole |
Hobbs, Horton H., Jr., and Henry W. Robison. A new crayfish of the genus Procam-
Por mmOonmEsolihnwestern Arkansas =.f.2u28.5552).. eee tel gee ee
Holt, Perry C. A new species of the genus Cambarincola (Clitellata: Branchiobdellidae)
from Illinois with remarks on the bursa of Cambarincola vitreus Ellis, 1919, and the
MRMISMONE NT OOKIMS AOI, L9G8. 0 ee
Huddleston, Richard W. Comments on the nomenclatural status of the families Cau-
cascliade and Favusellidae (Foraminiferida) .._...... =
Huddleston, Richard W., and Drew Haman. Nomenclatural status of the foraminiferal
MAMIE OOCITCIG SAIGON A 1980
Huddleston, Richard W., and Drew Haman. Jascottella, nom. nov. for Mamilla Scott,
1974 (Microproblematica) non Fabricius, 1823 (Mollusca) ____________________-----
Hutchings, P. A., and C. J. Glasby. Two new species of Ceratonereis (Polychaeta:
Nereididae) from estuarine areas of New South Wales, Australia ___-_
Kenk, Roman. Freshwater triclads (Turbellaria) of North America. XIII. Phagocata
RUMDLONAG Me Was pe cles, from sNeVaday, werd sie eee A
Kenk, Roman, and Anne M. Hampton. Freshwater triclads (Turbellaria) of North
America. XIV. Polycelis monticola, new species, from the Sierra Nevada range in
Pe EMOTE gee Selly ts ee eater yy ipl rl EEN RI coer) sarcu f wl ey inyrteesl bOI. ee oil

203-209
238-250
781-787

807-825

358-370

688-693

594-601

194-197

537-544

392-397

93-98

319-324
530-536

639-641
621-624

13-26

647-656

377-385

423-427

297-318

545-553

251-255

637-638

114-115

42]

515-521

161-166

567-570

Kensley, Brian, and Gary C. B. Poore. Anthurids from the Houtman Abrolhos Islands,
Western Australia (Crustacea: Isopoda: Anthuridae)__.----- =
Kornicker, Louis S. Alternochelata lizardensis, a new species of myodocopine ostra-
code from the Great Barrier Reef of Australia (Rutidermatidae) __._-
Kudenov, Jerry D. Redescription of the major spines of Polydora ligni Webster
(Polychaeta::Spionidae) 2-2 8tosat Vines wi ea ee ee Rn geeeaee le Od 2 eee
Kudenov, Jerry D., and John H. Dorsey. Astreptosyllis acrassiseta, a new genus and
species of the subfamily Eusyllinae (Polychaeta: Syllidae) from Australia _____________
Kyte, Michael A. Ophiacantha abyssa, new species, and Ophiophthalmus displasia
(Clark), a suggested new combination in the ophiuroid family Ophiacanthidae (Echi-
nodermata: Ophiuroidea) fromoff Oregon, (USA. 2 ee eee eee
Lewis, Julian J. A diagnosis of the Hobbsi group, with descriptions of Caecidotea
teresae, n. sp., and C. macropropoda Chase and Blair (Crustacea: Isopoda: Aselli-
aS) ona a a nN CI 2 A Sag SS Ao
Louton, Jerry A. A new species of Ophiogomphus (Insecta: Odonata: Gomphidae) from
thebwestern Highland Rit i Weminesse ec) See ae
Lynch, John D., and Pedro M. Ruiz-Carranza. A new genus and species of poison-dart
frog (Amphibia: Dendrobatidae) from the Andes of northern Colombia __________________
Manning, Raymond B., and Ch. Lewinsohn. Rissoides, a new genus of stomatopod
crustacean from the east Atlantic and South Atnica |= 22 ee eee
Manning, Raymond B., and Marjorie L. Reaka. Gonodactylus insularis, a new sto-
matopod crustacean from Enewetak Atoll, Pacific Ocean ___________________________-___-
Marshall, Harold G. Phytoplankton distribution along the eastern coast of the USA.
IV. Shelf waters between Cape Lookout, North Carolina, and Cape Canaveral,
Fikrig ee ee Rete eS aan er ae Ne ne
Martin, Joel W., and Lawrence G. Abele. Naushonia panamensis, new species (De-
capoda: Thalassinidea: Laomediidae) from the Pacific coast of Panama, with notes on
the Semis wih.24 2 es Tee ee See i ESE SORA (Ta aae Un he ee
McEachran, John D., and Janice D. Fechhelm. A new species of skate from Western
Australia with comments on the status of Pavoraja Whitley, 1939 (Chondrichthyes:
RaAjifOrmes 2 ttrts Sem Sn eee je 2 re et se RE es ae nD
McEachran, John D., and Janice D. Fechhelm. A new species of skate from the western
Indian Ocean, with comments on the status of Raja (Okamejei) (Elasmobranchii:
Rajiformes)i 2.2 20. es a od A Mea Ane oe a ees
McKaye, Kenneth R., and Catherine MacKenzie. Cyrtocara liemi, a previously un-
described paedophagous cichlid fish (Teleostei: Cichlidae) from Lake Malawi,
Atnica: 25 iieasee van! er he erie) ae iy Caer Be ee a

McKenzie, K.G. Description of a new cypridopsine genus (Crustacea: Ostracoda) from .

Campbell Island, with a key to the Cypridopsinae __________________________-__-__-
Mendez, G. Matilde, and Mary K. Wicksten. Notalpheus imarpe: a new genus and
species of snapping shrimp from western South America (Decapoda: Alpheidae) ______
Murdy, Edward O., and John D. McEachran. I[stigobius hoesi, a new gobiid fish from
Australia (Perciformes !Gobudae)i aie. 1 nn ee ee ee Gas eee ee
Nakamura, Izumi. Lateral line of Diplospinus multistriatus (Teleostei: Gempyli-
GAS) ORL. oie PE oh OE oP Oe IM We oie cel Sais, RI a A I
Nakamura, Koichiro, and C. Allan Child. Three new species of Pycnogonida from Sa-
gami'Bay / Japa. oie bal a
Opell, Brent D. A new Uloborus Latreille species from Argentina (Arachnida: Araneae:
MWPOD ORAS Ry haa OU eo ee cp
Parenti, Lynne R. Relationships of the African killifish genus Foerschichthys (Teleostei:
€yprinodontitormes:/Aplocherliad ac), Mase a ae eee,
Petuch, Edward J. Paraprovincialism: remnants of paleoprovincial boundaries in Recent
marie molluscan' provinces! ._c.ONiiieh Me i ie eee ae Se eee eee
Rausch, V. R., and R. L. Rausch. The karyotype of the Eurasian flying squirrel, Prero-
mys volans (L.), with a consideration of karyotypic and other distinctions in Glaucomys
spp? (Rodentia® SGnikidae) rl v* seer ee OL: AE A ee ee
Rohr, David M., and Richard W. Huddleston. Yochelsoniella, nom. nov., a new name
for Ellisella Rohr, 1980 (Gastropoda) non Gray, 1858 (Coelenterata) _____-_--___________

Vl

625-636

793-806

571-574

575-578

505-508

338-346

198-202

557-562

352-353

347-351

99-113

478-483

440-450

398-402

766-771

709-713

642-646

408-411

282-291

554-556

451-457

774-780

58-66

269

Rosewater, Joseph. A new species of the genus Echininus (Mollusca: Littorinidae:
Pehinmumae) with a review of the subtamily ==> 2 i
Schultz, George A. Amerigoniscus malheurensis, new species, from a cave in western
@reconm(@nustacea: Isopoda: Trichomiscidae)).. Bs... eee
Shelly, Roland M. A new xystodesmid milliped genus and three new species from the
eastern Blue Ridge Mountains of North Carolina (Polydesmida) _________________________-
Stauffer, Jay R., Jr., Brooks M. Burr, Charles H. Hocutt, and Robert E. Jen-
kins. Checklist of the fishes of the central and northern Applachian Mountains ______
Sterrer, Wolfgang, and Thomas M. Iliffe. Mesonerilla prospera, a new archiannelid
PROMI AnIMeNeAIVeStIN BEfMmmdar i096 ee
Thomas, Richard. A new dwarf Sphaerodactylus from Haiti (Lacertilia: Gekkoni-
Ty) eT NNR Ic ce 2 heh Td Oe eS ee ee es oe ied
Thompson, Fred G. A new species of Euglandina from Peru (Gastropoda: Pulmonata:
SHMUEL SCORE) a yl an al eR epee 0 ee
Thompson, Fred G., and Jane E. Deisler. A new tree snail, genus Drymaeus (Buli-
MUNA nOmMESOutMeastemm PTW 28
Uebelacker, Joan M. Review of some little known species of syllids (Annelida: Poly-
chaeta) described from the Gulf of Mexico and Caribbean by Hermann Augener in
I eran eaten erie a eter fect ee RT ed Po es te ee
Vari, Richard P. Hemiodopsis ocellata, anew hemiodontid characoid fish (Pisces: Char-
AcCHMeMetLoOmiwestern OUNIMAM oo se
Vari, Richard P. Curimatopsis myersi, a new curimatid characiform fish (Pisces: Char-
aciformes) from Paraguay scale), eM tars nee Gee Aemtee ah eee ROBO ON ee
Vecchione, Michael. Morphology and development of planktonic Lolliguncula brevis
eb a mClarm Vin OPSIGa)) a. 2 et ee ee ee
-Wainright, Sam C., and Thomas H. Perkins. Gymnodorvillea floridana, a new genus
and species of Dorvilleidae (Polychaeta) from southeastern Florida _____-___---__________
Wicksten, Mary K. New records of pinnotherid crabs from the Gulf of California
REM OuicmeIIMOUNSHIGAG) <2 82. 6 St eS ee
Wicksten, Mary K. Pinnixa costaricana, a new species of crab from Central America
REM nay em EenTIMOUNCTIGAC) =. ot etl eae Ne on ye ee ee
Zibrowius, Helmut, and Stephen D. Cairns. Remarks on the stylasterine fauna of the
West Indies, with a description of Stylaster antillarum, a new species from the Lesser
Paaiiilesnenmadania: Eydrozoa: Stylasterima) 20... 0
Zottoli, Robert. Two new genera of deep-sea polychaete worms of the family Amphar-
etidae and the role of one species in deep-sea ecosystems ___________________-----
Zusi, Richard L., and Gregory Dean Bentz. Variation in a muscle in hummingbirds and
MiSEAMGniseSyStematic implications 2.2222... 21.2222 eee een es

vil

458-477

27-47

509-514

81-88

763-765

265-268

583-593

188-193

788-792

602-609

694-701

354-357

579-582

210-221

48-57

412-420

INDEX TO NEW TAXA

VOLUME 95

(New taxa indicated in italics; new combinations designated n.c.)

COELENTERATA
Hydrozoa

Stvlaster antill arin sist eo aes Oe eR ee Mee

Callegorpia chariessa@..25 02's * i se ea ee Oe Se a tal
Fanelliaxcoryimbosa@ 20022 es oe ee

PLATYHELMINTHES
Trematoda

Atrophecaecum lobacetabulare wien 2 eat ee 2 eee

Phagocatashamiptonae 20235400) oe, ue a ies oe ee
Polycelis, montiCola; 20x 20s. ey. sed ope Dee ga esa pi SS el eel ee ea

ANNELIDA
Archiannelida

Mesonenilla prosper 25 aa SIE ae a
Cambarincola illimoisensis 122502 es ee ee ie ne

AiSORE DOSS se SE a ae Tae teal
CV ASSUSCLG [2.0 WNT. DRS DO Se I a Ss Ss ee Ore Ne

CELE VOWS ee ie hi este Dp Pl pe aE a ea
Decemunciger’ ia. \ S002 aie be EO SS Se las Oe eee ne ne
GPQNEGW Ec Fly WN RT Re CSR Re ahs a i cele A te ga real
Brdecamera. 02:02. Rae. NU Ce oc ON eg) EAR Re 2 A eee
Paleg 'x.weooy ee TY AYE SE: SAN ae Be 1 ad
Eunice: pulvinopalpata. 2040) 22 PS 2 he RR on he ES Ee Se reece a
GyMVWOU OPV CO a LEO ES AA ce ets) cea lied Jo eee oe len eg
flonidang) Ga... LSA) EOE SUA of EE AR ea So a ee
Haplosyilis floridamat.C.. fe eg ce
Mooreonuphisyonesi 2000. 8 Re ee SIDE My SN Sept eee nd gd
NG@tOmaStuls (Cai eHi oes os ise ae I a a a

Onuphiss(Nothnia)*australatlantica aa ee eee ee

heterodentata. 22. es a ae Ee

lithobiformis ¢ 2 to 2 0es) DS aie ie eM aaa Rae Ae ae

(Onuphis) declivorum 3204 ee ee ee ee eee

LiffiCilis sc eT) Ce a EINE ULES Tio ial te en aren LO ee ee

OV CWS QUIUZE 9 Sa he ee a we Pee | el i cis eh ee a

D011 ce Ee A TEA COW TM NL at MeN Me Meee NG aS Se A

Pavalvinehl@:, eo 0 fe ks i See le 5 TE ae (Te Pe en na ee
GU GIST A. SRN ata Nera Iie A led Pc I sg aD
UNCISPIONIDAE =) Mik ceck) Oe EO Ree ce Ies CR aa ee A A ee eS See

TWEE SIDUD ence telco eee al I IR SOP eA a) SE ae ce RRS DEL aes a SO 534

[AO TTGUUOID gee le TE lat a Tee TES a7 i ATI aR oO ot ne MY SRR SE ES 534
ARTHROPODA
Crustacea

Meet Onen Se ARCMICCL CLETUS IS pee wate sche aos ver ne OW et eS ke a ee 793
PRIME TO OMS CUISMOULCHNCNSIS) waste ot Sie Bere Oy SN ee 89
ELSIE, CPA TACNT LeNS aaa ie ce t d SE SAAAn e niece Sy?)
SCVMM LOCUPTO pe SO a PT me lia al Re SS et PSP en RO pe een eo eM 625
PAW ae) TCAEET CA) (0 ene TE Ge) on) Vu hee ee a Ne NU A oe iy ee Ne 171
ALAR D CDV TIOID SES sg ge a 8 SIE RI Re mR gi Se, a a a el 766

ROTA ce oe seeps Se pa OT cle ae A ek LT, Aveta oe ee OPE 768
BAA CLOTS IACI Camm mee a tic Re Ng ee I Ss as ES pe ge i Lo Oe RIL | Pv ge 339
WORE OL OLACMSUS 52 ee eee aig tne Renee nen re, Re ee PL se nee 538
Peer cnter MSIE SCI OSAMRI cae os Ne Ee A a a 319
OE GIEE DM CHCYB OS aks = Ae EEN TS bal SE i Ae lepetet Ear DNIeN ernee iee Eee 703
ESR HVE CS SMO CUACAD 1G Comer nme eee rnin 2 eet er Oy ie Ol a AAA OL ee pe nt Pode tsar 628
EO AID TION MCOMIID) AU ee ae ee I I Ee 610
LEENA TLIO! is ET are Pe 8 a le Se ne ee 182
SSAA CICS SLL SS A NS er ws Ne 08 Sb ea np ea Wee 347
PREM OSOMMAB IIL DUNEMSIS: Bi. 2 Vl a) 20 yet bie a 547
eer (ares) Fr eRe Eh ae) ee ahr ra) uh hele LO leet i i ea A ea as 495

NOUDISCUIS epee Sie 2 SRO NS ES Oe a oe eee eee ee ete Oe RE mS I nee 495

rea | | ee onl weit Ne pee SP MNES Ae BE ee a eS So Eas 502
EEE OU SADE METS, cena an A Bg AR SE PD 180
AWHONN ODE a eecsecien ae eS ep a YS PRT ctl le UDO ened Ad YS pk 182
erent RULER anh CLC EE een aS, ee PN NTE AD foes i age a ee ee 540
CLRPURGOVEOYET I Ba a ay Ne eer Deh BA Oe a REE Le ee pec ee aU 174

eh Coenen eR eS Ya eh Of Re uees Wl elie Wires tS eh eee ae Se 174
Re et) 7 nC Ree en mevmetn ns we ee My Sha iE re eS ete 176
MARLON OMIR ERI CHIOC) GC aemmne elmer ct Loe! eee ee 8g os se totes Sh NS ee 321
IE ERE VE ee tik Bs A sl a a Ee a ia en VP 170
fla MuSlivCUMD: (PUPUAILTAUCIASIIS cick Se NAA Sy BS Teg ar De pee NN te eal oe em ger ee ee ee 478
POSES CLERIG ASPROWIDE tec, sl tr ae Se ese ug esc ete ce ar aA OR cee IL a oe hE 709
ane easel eile Cah O)/) Spee ween me ee ye LE Me RE pels et ies eS ee 542
Sp SM SEGINTERE EPICUGW OND: 2 I TN I te ee eS 634
AUR COAL IESE MIT YAU S10 ne eT ln a ee Ee Sa 181
Me TINO CMC MSE MIM: Coree sep ew Ac See tt oe he ee ON ee Ne 170
MIRIAM TOR ST GAIT Lm ee ee el elles bam WE Nita be! OE ee le oe 579
EN eAMoanisn(Gimandiella) parasimulans 0 545
SABIE DERDCDEL AED, a dapat Sh Spee le eR D0 SS nce ae 168
ESN CHEGUEIS aan ancn cnet og ck lt A ce Pe ed, ERIN ete 352
ae ar Rte Tem Re A ei ee Vine on ie oa kN ee RT 169
MRC BGIER TCIM ANS LCLIIT LC eames Re Deon a eg ED oe fo a ei oe 93
IVI oo I ce a) a A ee pe I GE De es 176
fees BCU ELOISE ls ys tr cn ee eee eter one ERA 7)
ee oc | ee eee Je RT Pgs ona bs ho es es le 184

Insecta
ARTO MMOMUISH DOUGH CRG) (seeee ents! (2) ee ess ee ee ele oe a ah Pe 198
Myriapoda

EW OMGILI. pada SE aa ee a Rg SE AR a, 460

(AAA OTIS Seve Soe LA £9 ST NR SSE a we OIRO he SEN le BP 2) pe TN ONE AE Wes Cen ene hs 467

COMET OUUS! es a oS re 3 AOS IIT © PSE ep Neg ar Re Oe Os 463

fil RADIUS a 5 ps a NOT LUD LW NT NON nd Pc 0s yo ae eR ee ap See eee 470

Pycnogonida

Ammiothella:stawroniata 2 2.5 ce A Oe eae, 2 a ee eR ae eae kT RS, ee ae Dg
Anoplodactylus wiarsiiall crisis’): 225. els sacs ale EG ues Usps etalon a er Oe etek 274
perforatus 200) so Bet aE Ei ane ne rs ee 289
SRUIODGEMSIS( co 5) AN a8 SO ASS TO SNe ep eg LA dR leg oe 285
ACS COVEY ICIS Lette OUIaEE $y 00 i ek WN AE ala a eG ig ol oh oe 283
Nymphon microne sicuni cee 6. 08 2a eer ets Tie ae ES URE ne ns ree ey 27
Arachnida
Uloborus elongatus [Leite oes lye ti wena ea a I | a ete me ie 554
MOLLUSCA
DrymMacus aurantiOstomus: 28 Bs ent Es PEN Ae ee eta Ces 265
Echinanus vivipariis 2.002) 20 ec) A ie SSN ee ee kee ee 69
Euplandinay easy | occ es ee SS a re 763
Oreohelix ‘litoralis 20.220 et ce a hE rte is Mee ars ye 256
Pterynotus (Pterynotus) xenwos” loc eat Le a 639
VOCHE] Sore dha 0500 ae ah oR eR IT Ha Lor Te Wate tt reper ea 269
ECHINODERMATA
Evoplosomia: virgo ‘stot suet tec 6th SER is eee ea 0 Re Pole a en eo Te,
Ophiacanthavaby sear 42 1k tk ee ee ee ee ee 505
Ophiophthalmus diplasia ac oe er ee ee ee 508
CHORDATA
Pisces
Archomentdiatwvarvel@e: 25! oe ag NT he Tre ea 430
GurimatopsisS miyerst Loo ee RE ee a 788
Gyrotocara dient < on. oe 398
Hemiodopsis ocellata (cs 2 tee a es 188
Hy pselognathus horridus | 2-22. jus 403
ISHPODIUS HOST 6 oe re es ce eae 643
Micro pnathuls: natans. 2068 el ae Mio ee ea te 682
Bay onayanallenigw qn. 2 2s 2 te ee NE So de eae 8
Rajan(Okamejel)eemstrat =... e 44]
Amphibia
AM OD OPTATVTUES aa il a NR IB RS > er 57)
SVILOMMOPUS) chest a LN Ra ae A Ry le ec 0 ek Ai an a DSI)
Piylodes baba@x tse hs ie 380
WOUTUZOLEIUTT Dees oo OA ue say 3 Tea 9 Onan aed Lt eg IT ee 382
Platyamantisispelaeuis: 26 o6 0.0 2 ts os, oi Ue A age I eS 386
Reptilia
Sphaerodactylusseelicana.. oto 2 ss Se 392
VILO CV UO ep I alo Se we RN pS SIRI A SR 81
Mammalia
Podogymmura: aureospinila@ a5 oo I Ta A a Sn ae ca oe 14
Sasuinus mignicollis*hernandezi. 25.2) a ne 649
MICROPROBLEMATICA
TGS COPLSN 05 5h MI he Bah gO el A ie a Sc aR 421

S74, 0618

(ISSN 0006-324X)

Proceedings
of the
BIOLOGICAL SOCIETY
of
WASHINGTON

F QAO NIAA
Libh AM Kt

Volume 95 13 April 1982 Number 1
;

THE BIOLOGICAL SOCIETY OF WASHINGTON

1981-1982
Officers

President: Raymond B. Manning Secretary: Michael A. Bogan
Vice President: Paul J. Spangler Treasurer: Leslie W. Knapp

Elected Council

Frederick M. Bayer Isabel C. Canet
Kristian Fauchald David L. Pawson
Austin B. Williams

Custodian of Publications: Michael J. Sweeney

PROCEEDINGS
Editor: Brian Kensley Co-editor: Stephen D. Cairns

Associate Editors

Classical Languages: George C. Steyskal Invertebrates: Thomas E. Bowman
Plants: David B. Lellinger Vertebrates: Richard Banks

Insects: Robert D. Gordon

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Second class postage paid at Washington, D.C., and additional mailing office.

PROC. BIOL. SOC. WASH.
95(1), 1982, pp. 1-12

A NEW SPECIES OF SKATE FROM WESTERN
AUSTRALIA WITH COMMENTS ON THE
STATUS OF PAVORAJA WHITLEY, 1939

(CHONDRICHTHYES: RAJIFORMES)

John D. McEachran and Janice D. Fechhelm

Abstract.—The genus Pavoraja is resurrected for the Australian skates Raja
nitida and Pavoraja alleni, n. sp., which differ from all recognized genera of
skates in clasper and rostral structure, and by their combination of derived char-
acter states. Pavoraja alleni is described and diagnostic characters are given for
Pavoraja and P. nitida.

Raja nitida, which occurs off Tasmania, Victoria, and southern New South
Wales (Whitley 1940), was placed in a new genus Pavoraja, along with another
_ eastern Australian skate, Raja polyommata Ogilby, 1916 by Whitley (1939). Whit-
ley (1939) only briefly described Pavoraja and gave no characteristics that dis-
tinguish it from other skate genera. The following year Whitley (1940) offered
little to augment his description, but stated that P. nitida resembles the South
American skate, Malacorhina scobina (=Psammobatis scobina Philippi, 1857),
that the type locality of P. nitida may be South America rather than Australia,
and that the Australian specimen of P. nitida may represent a new species or
subspecies. More recent authors (Fowler 1941, Bigelow and Schroeder 1953)
considered Pavoraja synonymous with the genus Raja, because of Whitley’s lack
of a generic diagnosis.

During investigations of the skate fauna of Western Australia a new species
was discovered which closely resembles Raja nitida Gunther, 1880. Herein we
describe the taxonomically important anatomical characters of R. nitida, com-
ment on the taxonomic status of Pavoraja, describe the new species, and then
comment on the relationship of R. polyommata to the other species.

Materials and Methods

Specimens of Raja nitida, R. polyommata, and the new species were obtained
from the Australian Museum, Sydney (AMS); British Museum (Natural History)
(BMNH); Tasmanian Museum, Hobart (TMH); and Western Australian Museum,
Perth (WAM). Several individuals of R. nitida and one of the three specimens of
the undescribed species were dissected to reveal the structure of the claspers,
neurocrania, and scapulocoracoids. Several additional specimens of R. nitida and
all three specimens of the new species were radiographed to verify the anatomical
observations based on dissections and to count vertebrae, pectoral radials, and
pelvic radials. Methods for making measurements and counts are described by
McEachran and Compagno (1979, 1982).

Z PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1Icm

sl
cf

SI.

a b

Fig. 1. Lateral view of right clasper, partially expanded to show components: a, R. nitida, AMS
1B.5275; b, Pavoraja alleni, WAM P19118 (Holotype). cf—cleft, hy—hypopyle, pr—pseudorhipidion,
rh—rhipidion, sl—slit, sp—spike, sr—spur, st—sentinel.

Results

Claspers.—Raja nitida has very slender, short claspers which are constricted
rather than expanded at origin of glans (Fig. la); without dermal denticles or
pseudosiphon; inner dorsal lobe with pseudorhipidion extending from level of
hypopyle to about distal one-third of glans, continuing distally as a fold of integ-
ument; slit located lateral to pseudorhipidion; spur well developed; cleft medial
to spur; rhipidion well developed, running from level of hypopyle to distal one-
third of glans, distal section rotated laterally and running over base of sentinel;
sentinel rod-shaped and covered with integument, extending from level of slit to
near tip of glans; spike ventral to sentinel, located within sentina, disc-shaped
with a sharp, naked lateral margin; axial cartilage forming a slender tip (Fig. 2a,
b, c); dorsal marginal little expanded distally, distal margin truncate, with an
inner distal extension entering glans and forming pseudorhidion; ventral marginal
with a evenly convex distal margin; dorsal terminal 1 and ventral terminal mem-
branous, broadly joined on ventral aspect of glans, forming a sheath-like covering
of glans; dorsal terminal 2 broad, fused to distal and distolateral surface of dorsal
marginal; dorsal terminal 3 fused with dorsal terminal 2, small with a distally
pointed and laterally curved extension forming spur; ventral terminal V-shaped,
lacking a sharp lateral margin, lying on dorsal surface of accessory terminal 1;
accessory terminal 1 Y-shaped, fused with distal surface of ventral marginal,
S-shaped distal extension forming sentinel; accessory terminal 2 short, attached
to accessory terminal 1, with a dorsoventrally flattened, disc-shaped extension
forming spike.

VOLUME 95, NUMBER 1 3

Fig. 2a, b, c. Right clasper cartilages of R. nitida, AMS 1B5275: a, Lateral view, partially ex-
panded with dorsal terminal and ventral terminal cartilages shown separately; b, Dorsal view; c,
Ventral view; d, e, f, right clasper cartilages of P. alleni, WAM P19118 (Holotype), d, Lateral view,
partially expanded with dorsal terminal and ventral terminal cartilages shown separately; e, Dorsal
view; f, Ventral view. Atr,—accessory terminal |, atr.—accessory terminal 2, ax—axial, dmg—dorsal
marginal, dtr,—dorsal terminal 1, dtr,—dorsal terminal 2, dtr,—dorsal terminal 3, vmg—ventral mar-
ginal, vtr—ventral terminal.

Neurocranium.—The neurocranium of R. nitida has a short, slender rostral
Shaft, fused to flattened rostral node and appendices at tip of snout (Fig. 3a),
widely separated from rostral base and neurocranium; rostral appendices free of
rostral shaft posteriorly and possessing two foramina; propterygia of pectoral
girdle reaching rostral appendices; nasal capsules of moderate size, set at about
a 40° angle to transverse axis of neurocranium; foramen for profundus nerve on
leading edge of nasal capsule; anterior foramen for preorbital canal on anterior
margin of kidney-shaped basal fenestra of nasal capsule; precerebral space nar-
row, inner walls of nasal capsules moderately bulging into precerebral space;
interorbital region narrow (Table 1); preorbital processes poorly developed, sep-
arated from supraorbital crests by shallow notch; postorbital processes well de-
veloped; anterior fontanelle tear-shaped; posterior fontanelle trapezoid-shaped;

+ PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Neurocranium of R. nitida, AMS 1B.5275: a, Dorsal view; b, Lateral view; c, Posterior
view; d, Ventral view. ac—anterior cerebral vein foramen, af—anterior fontanelle, antc—antorbital
condyle, bf—basal fenestra, end—endolymphatic foramen, es—efferent spiracular artery foramen,
hf—hyomandibular facet, ic—internal carotid artery foramen, into—intercerebral vein foramen, ja—
jugal arch, ibX—lateralis branch of vagus nerve foramen, nc—nasal capsule, obf—otic branch of
facial nerve foramen, of—orbital fissure, onc—orbitonasal canal, os—optic stalk, peri—perilymphatic
foramen, pf—posterior fontanelle, poc—preorbital canal foramen, postp—postorbital process, prep—
preorbital process, prof—profundus nerve foramen, ra—rostral appendix, rb—rostral base, rn—ros-
tral node, rs—rostral shaft, I—optic nerve foramen, I1J—oculomotor nerve foramen, 1!V—trochlear
nerve foramen, VIJ—hyomandibular branch of facial nerve foramen, [X—glossopharyngeal nerve
foramen, X—vagus nerve foramen.

foramen for anterior cerebral vein posterior to line connecting foramina for preor-
bital and orbitonasal canals (Fig. 3b); trochlear nerve foramen posterior and dor-
sal to optic nerve foramen; oculomotor nerve foramen situated above optic stock;
orbital fissure located on posterior aspect of orbit, anterior to foramen for hyo-
mandibular branch of facial nerve and posterior to foramen for interorbital vein;
efferent spiracular artery foramen on ventral rim of orbit; jugal arches moderately
slender; vagus nerve foramen immediately ventral to foramen for lateralis branch
of vagus nerve and medial to foramen for glossopharyngeal nerve (Fig. 3c); basal
plate moderately narrow (Fig. 3d).

Scapulocoracoid.—The scapulocoracoids of R. nitida are moderately short and
anteroposteriorly elongated (Table 2), without an anterior bridge (Fig. 5a); an-
terior fenestra little expanded; postdorsal fenestra moderately expanded; meso-
condyle not expanded; three postventral foramina, first greatly enlarged; neo-
pterygial ridge between mesocondyle and metacondyle incomplete.

Comments.—Raja nitida differs from all other species of Raja in clasper and
rostral structure and differs from all other genera of skates in its combination of
derived character states, suggesting that the genus Pavoraja should be resur-
rected for this species. ‘‘Raja’’ nitida differs from Raja in possessing claspers
with membranous dorsal terminal 1 and ventral terminal cartilages which are
broadly joined along the ventral aspect of the glans and disc-like accessory ter-
minal 2 cartilages; a reduced and incomplete rostral shaft which is widely sepa-

VOLUME 95, NUMBER 1

Fig. 4. Neurocranium of P. alleni, WAM P19117 (Paratype)
Posterior view; d, Ventral view. Abbreviations as in Fig. 3.

“CTS
a.

: a, Dorsal view; b, Lateral view; c,

rated from the rostral base, and propterygia of pectoral fins which reach rostral
appendices. Reduction of the rostral base precludes this species from being clas-
sified in Raja (Ishiyama 1958, Stehmann 1970, Hulley 1972, McEachran and Com-

pagno 1982).

Table 1.—Neurocranial measurements of P. alleni and P. nitida expressed as percentage of na-

sobasal length.

Nasobasal length (mm)
Cranial length

Rostral cartilage length
Prefontanelle length
Cranial width

Interorbital width

Rostral base

Anterior fontanelle length
Anterior fontanelle width
Posterior fontanelle length
Posterior fontanelle width
Rostral appendix length
Rostral appendix width
Rostral cleft length
Cranial height

Width across otic capsules
Least width of basal plate
Greatest width of nasal capsule
Internasal width

P. alleni
278 mm TL @

26.7

185
23
84
32
26
13
33)
14
42
Il 2
16
15

6
24
58
24
37)
13

P. nitida
335 mm TL ¢

36.2

168
20
70
80
al
10
29

9
47
17
14
10

6
23
50
25
37
10

P. nitida
357 mm TL 3

40.1
Sil
24
70
80
ns)
Ih
28
8
34
11
IU
1]
>)
22
Sil
24
Sy
11

6 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Scapulocoracoid measurements of P. alleni and P. nitida expressed as percentage of
scapulocoracoid length.

P. alleni P. nitida P. nitida
278 mm TL ? 335 mm TL 3 357 mm TL 3
Scapulocoracoid length (mm) 16.3 Ds Dll
Scapulocoracoid height 71 74 I)
Premesocondyle 39 40 41
Postmesocondyle 61 60 58
Postdorsal fenestra length 32 3 33
Postdorsal fenestra height 24 WS) 19
Anterior fenestra length V7 15 1S)
Anterior fenestra height DS 24 Dy
Rear corner 53 59 58

Pavoraja Whitley, 1939

Type species.—Raja nitida Gunther, 1880.

Diagnosis.—Snout with small laterally compressed rostral process; caudal fin
poorly developed, with an epichordal lobe and with or without a hypochordal
lobe; claspers constricted at origin of glans; dorsal marginal with a distomedial
extension forming pseudorhipidion; dorsal terminal 1 and ventral terminal mem-
branous and broadly joined along ventral aspect of glans; dorsal terminal 3 with
free distal tip forming spur; accessory terminal | with an S-shaped distal extension
forming sentinel; accessory terminal 2 with a disc-shaped extension forming
spike; rostral shaft slender and reduced, separated by a wide distance from nar-
row rostral base; rostral appendices short, propterygia of pectoral girdle extending
to rostral appendices; nasal capsules with basal fenestrae; anterior fontanelle and
internarial plate moderately narrow; interorbital region narrow; preorbital pro-
cesses poorly developed; foramen for anterior cerebral vein posterior to line

scp

pdfe

mtc

msc
b
pvf
Fig. 5. Lateral view of scapulocoracoid: a, R. nitida, AMS 1B.8274; b, Pavoraja alleni, WAM

19117 (Paratype). af—anterior fontanelle, msc—mesocondyle, mtc—metacondyle, pdfe—postdorsal
fenestra, prc—procondyle, pvf—postventral foramina, scp—scapular process.

VOLUME 95, NUMBER 1 7

connecting foramina for precerebral and orbitonasal canals; scapulocoracoid sub-
rectangular to almost rectangular, moderately short and anteroposteriorly ex-
panded; without anterior bridge; three postventral foramina; precaudal mono-
spondylous vertebrae ranging from 26 to 29 and predorsal caudal diplospondylous
vertebrae ranging from 66 to 79.

Remarks.—Arhynchobatis, Bathyraja, Breviraja, Gurgesiella, Psammobatis,
Pseudoraja, “‘Raja’’ waitei, and Sympterygia also possess reduced rostra, a de-
rived state (McEachran and Compagno 1979, 1982). Rostra of Arhynchobaitis,
Bathyraja (in part), Psammobatis, ‘‘Raja’’ waitei, and Sympterygia are basally
segmented and could presumably have evolved into the P. nitida state by distal
retraction of the rostral shaft. The rostral shaft in Psammobatis 1s partially re-
tracted while that of Pseudoraja is nearly absent (McEachran and Compagno
1979). However, all of these taxa differ from P. nitida in structure of the claspers,
neurocrania and scapulocoracoids (Compagno and McEachran, in prep.) and thus
it seems likely that rostral reduction has occurred separately several times within
the skates. Pavoraja nitida shows a closer relationship to the genera with reduced
but basally unsegmented rostra (Breviraja and Gurgesiella) in structure of the
neurocranium and scapulocoracoid, and to a lesser degree, in clasper structure,
and possibly could have been derived from a common ancestor of either of these
taxa. Breviraja possesses a distally segmented rostral shaft which could have
_ evolved into the P. nitida state by basal retraction of the proximal, unsegmented
part of the shaft. However, uniqueness of the clasper and rostral structure of P.
nitida precludes its classification with either Breviraja or Gurgesiella and sup-
ports the resurrection of Pavoraja.

Based on the examination of several small immature specimens of R. polym-
mata, this species is classified in Raja rather than Pavoraja. It possesses a stout
rostral shaft and in all aspects agrees with the anatomical character states within
Raja. Further comments on its relationships must await the procurement of large,
mature specimens to make more extensive anatomical observations and to ex-
amine the claspers.

Pavoraja nitida (Gunther, 1880)
(Fig. 6, Table 3)

Diagnosis.—Disc width greater than 54% of total length; orbital length 1.7 to
2.1 times as long as spiracle; anterior margin of pelvic fins less than 75% of
distance from origin of anterior lobe to extreme posterior margin of fin; second
dorsal fin and epichordal lobe of caudal fin confluent; epichordal lobe shorter than
base of second dorsal fin; hypochordal lobe of caudal fin absent; dorsal surface
dark brown with small light spots, some of which are arranged into occelli; mature
males range from 312 to 363 mm TL.

Comments.—It is very unlikely that the type locality of P. nitida is South
America rather than Australia, as suggested by Whitley (1940). The holotype of
P. nitida does not differ significantly from the specimens from Australia. No
Specimens resembling P. nitida have been reported from South America, and
Specimens resembling this species have not been discovered in the large samples
of South American skates studied by McEachran.

Material examined.—BMNH 1879.5.14.417 (Holotype), Two Fold Bay, New

8 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Pavoraja nitida, BMNH 1879.5.14.417 (Holotype).

South Wales, Australia, HMS Challenger, 6 April4 June 1874. AMS 1A.2493
(1), AMS 1A.3904.05 (1), AMS 1B.4324.25 (2), AMS 1B.5275 (2), AMS 1B.8274
(3), AMS E.2165 (1), AMS E.5453 (1), AMS I.10823.24 (2), AMS I.16564.001 (1),
AMS 1A.3904.05 (1), TMH D816 (1).

Pavoraja alleni, new species

Holotype.—WAM P19118, 297 mm TL, mature male, collected off northwest-
ern Australia, near Rowley Shoals, 17°17.0’S, 119°57.0’E, 350 m, 20 December
1969.

Paratypes.—WAM P19117, 278 mm TL, female, collected with holotype;
WAM P8226, 159 mm TL, female, collected December 1963 or January 1964 in
the eastern Indian Ocean, 17°05’S, 119°48’E, aboard the Umitaka Maru.

Diagnosis.—Disc width less than 52% of total length; orbital length 2.2 to 3.2
times as long as spiracle; anterior margin of pelvic fins 81 to 104% of distance
from origin of anterior lobe of fin to extreme posterior margin of fin; second
dorsal fin and epichordal lobe of caudal fin not confluent, distance from posterior
margin of base of second dorsal fin to tip of tail considerably greater than length
of base of second dorsal fin; hypochordal lobe of caudal fin small but present;
dorsal surface light tan with minute dark spots loosely concentrated into ill de-
fined blotches but not forming ocelli; males mature at 297 mm TL.

Description.—Disc 1.1 times as broad as long; maximum angle in front of
spiracles 107° in holotype (110° to 112° in paratypes); margin of disc convex
except concave opposite orbits and spiracles; outer corners of disc broadly round-

VOLUME 95, NUMBER 1 9

Table 3.—Proportional measurements and meristic values for P. alleni and P. nitida. Pro-
portions are expressed as percentage of total length.

P.alleni P.alleni P.alleni P.alleni PP. nitida P. nitida P. nitida

(holotype) (paratype) (paratype) xy (holotype) n= 15 x
Sex 3 © 2 3
Total length (mm) 297 278 159 209 242-358
Disc width 50.8 49.3 47.2. 49 59 54-62 59
Disc length 44.4 43.5 42.8 44 50 45-52 Si
Snout length (preocular) 10.4 9.6 Ea eles 11.0 8.3-11.4 5)
Snout length (preoral) 9.8 10.5 22s O57/ 11.8 9.3-12.6 10.6
Snout to maximum width 28.6 26.3 BY 2 29 25-31 28.7
Prenasal length 8.1 8.0 Oe 8.4 8.2 6.3-9.1 Teil
Orbit diameter 4.7 3) 4.7 4.4 al 5.2-6.1 D5
Distance between orbits Doll Spl 3.3 3.0 31,2 2923.7, 353)
Orbit and spiracle length 325 4.6 D2 5.0 6.3 6.2-8.9 6.6
Spiracle length LS) 1.8 1.4 1.6 B)522 2.7-6.3 Shy?
Distance between spiracles 6.6 6.0 6.4 6.3 6.5 4.2-6.7 6.3
Mouth width 6.9 6.1 5.8 6.3 6.3 5.9-7.9 6.9
Nare to mouth SD) 3.5 4.5 oll gi) 3.24.7 4.2
Distance between nostrils 4.8 4.5 ayy 4.8 4.1 3.24.7 308)
Width of first gill opening 0.9 1.0 0.8 0.9 7 1.6-2.1 ow
Width of third gill opening 1.0 lest 0.8 1.0 159 1.5—2.0 lew
Width of fifth gill opening 0.7 0.9 0.8 0.8 1.6 1.3-1.6 1.4
_ Distance between first gill openings 12 12 2 2 13 15-12 12
Distance between fifth gill openings 8 8 a 8 7 6-8 6
Length of anterior pelvic lobe 12 11 13 12 12 12-14 13
Length of posterior pelvic lobe 13 14 13 14 16 17-18 17
Distance—snout to cloaca 41 4] 40 41 44 41-45 43
Distance—cloaca to Ist dorsal fin 47 47 48 47 46 43-47 45
Distance—cloaca to caudal origin 55 57 5) 35) 54 51-56 53
Distance—cloaca to caudal tip 2) 59 61 60 a7 54-59 57
Number of tooth rows (upper jaw) 38 359) 45 39 34 31-36 3)
Sample size of radiographs 3 4
Number of trunk vertebrae 26 28 26 26.7 ped 26-29 ies
Number of predorsal caudal vertebrae 79 71 74 74.6 66 66-72 70.2
Number of pectoral radials 63 67 65.5 WZ 62-73 69.8
Number of pelvic radials 16 15 IS).5) 20 19-20 IF)

ed. Tip of snout with small, laterally flattened process. Axis of greatest width
76% (67 to 68%) of distance from tip of snout to axil to pectoral fins. Pelvic fins
deeply incised, anterior lobe narrow and acutely pointed; anterior margin 85%
(81 to 104%) as long as distance from origin of anterior lobe to posterior extreme
of fin. Tail slender, little depressed, its width at midlength about two-thirds di-
ameter of eye. Tail with narrow lateral fold along ventrolateral surface running
from near tip of pelvic fins to origin of hypochordal lobe of caudal fin, widening
near tip of tail fold to equal height of epichordal lobe of caudal fin. Length of tail
from center of cloaca to distal tip 1.4 times (1.5 times) distance from tip of snout
of center of cloaca.

Preocular length 2.5 times (2.2 to 2.5 times) as long as orbit; preoral length 2.0
times (2.4 to 2.5 times) internarial distance. Interorbital distance 0.6 (0.7 to 0.8)
times length of orbit, orbit length 3.1 (2.2 to 3.3) times as long as spiracles.

10 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 7. Pavoraja alleni, WAM P19118 (Holotype): a, Dorsal view; b, Ventral view; c, Ventral
view of head.

Fig. 8. Pavoraja alleni, WAM P19117 (Paratype).

VOLUME 95, NUMBER 1 11

Anterior and posterior nasal flaps without fringes; without nasal pits. Upper and
lower jaws moderately arched (little arched). Teeth with pointed cusps near sym-
physes of jaws, with rounded cusps near margin of jaws (with rounded cusps
throughout jaws), teeth arranged in more or less transverse series (in quincunx).

Distance between first gill slits 2.5 (2.4 to 2.8) times as great as between nares;
distance between fifth gill slits 1.6 (1.3 to 1.8) times as great as between nares;
length of first gill slits 1.4 (1.1 to 1.2) times length of fifth gill slits and 0.1 (0.1 to
0.2) times mouth width. First dorsal fin slightly higher and longer than second;
interspace between dorsal fins equal to or slightly shorter than base of first dorsal
fin; second dorsal fin separated from epichordal caudal-fin lobe by distance equal
to one-half base of second dorsal fin; epichordal lobe low, length of base about
equal to that of second dorsal fin; distance from end of base of second dorsal to
tip of tail considerably greater than length of base of second dorsal fin; hypo-
chordal caudal lobe small.

Upper surface of disc, pelvics and tail densely covered with denticles. Ventral
surface naked. Orbit with 3 thorns on anteromedial margin and 3 thorns on pos-
teromedial margin; 3 prenuchal and 1 nuchal thorns; 3 irregular rows of thorns
on dorsal surface of tail, no interdorsal thorns. Holotype with 5 alar spines on
left and 4 on right lateral aspect of disc and a patch of malar thorns on antero-
lateral aspect of disc.

_ The claspers of P. alleni are similar to those of P. nitida with the following

exceptions: claspers very slender; rhipidion poorly developed; sentinel curved
laterally and extending to tip of glans (Fig. 1b); dorsal terminal | only loosely
connected to ventral terminal along ventral aspect of glans; dorsal terminal 3 with
a longer distal extension forming spur (Fig. 2d, e, f); ventral marginal with a
truncated distal margin; accessory terminal | with a relatively longer and less
curved distal extension forming sentinel; accessory terminal 2 with a more elon-
gated shaft supporting disc-like extension forming spike.

The neurocranium of P. alleni is similar to that of P. nitida with the following
exceptions: nasal capsules set at about a 30° angle to transverse axis of neuro-
cranium (Fig. 4); rostral base better developed, extending nearly to leading edge
of nasal capsules; precerebral space broader (Table 1), inner walls of nasal cap-
sules not appreciably bulging into precerebral space; postorbital processes poorly
developed; jugal arches relatively slender.

The scapulocoracoids of P. alleni are similar to those of P. nitida (Table 2)
with the following exceptions: the anterior vertical margin is more perpendicular
to horizontal axis (Fig. 5b); posterior corner more posteriorly located; postdorsal
fenestra is oval-shaped rather than elliptical.

Color.—Dorsal surface uniformly light tan with minute dark spots loosely con-
centrated into ill defined, symmetrically arranged blotches; tail darker with four
obscure brown bands; dorsal fins tan with brown blotches. Ventral surface light
tan.

Etymology.—Named after Gerald R. Allen (WAM) who furnished us with the
specimens of the new species.

Acknowledgments

We wish to thank Alwyne Wheeler and Peter J. P. Whitehead for providing
work space at the British Museum (Natural History); Gerald R. Allen (WAM),

12 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

A. P. Andrews (TMH), and Doug Hoese (AMS) for providing specimens; and
Matthias Stehmann for providing a radiograph and photograph of the holoype of
Pavoraja nitida. Figures la, 2a, b, c and 3 were prepared by Debbie Allen, other
figures by Janice D. Fechhelm. Helen Feney labelled and mounted the figures.

Literature Cited

Bigelow, H. B., and W. C. Schroeder. 1953. Fishes of the western North Atlantic, Part II. Sawfishes,
guitarfishes, skates and rays, and chimaeroids.—Memoir Sears Foundation for Marine Re-
search 1(2):588.

Fowler, H. W. 1941. The fishes of the groups Elasmobranchii, Holocephali, Isospondyli, and Ostar-
ophysi obtained by the United States Bureau of Fisheries Steamer “‘Albatross’’ in 1907 to
1910, chiefly in the Philippine Islands and adjacent seas.—United States National Museum
Bulletin 100(13): 1-879.

Hulley, P. A. 1972. The origin, interrelationships and distribution of southern African Rajidae (Chon-
drichthyes, Batoidei).—Annals of the South African Museum 60: 1-103.

Ishiyama, R. 1958. Studies on the rajid fishes (Rajidae) found in the waters around Japan.—Journal
Shimonoseki College of Fisheries 7: 1-394.

McEachran, J. D., and L. J. V. Compagno. 1979. A further description of Gurgesiella furvescens

with comments on the interrelationships of Gurgesiellidae and Pseudorajidae (Pisces, Rajoid-

e1).—Bulletin of Marine Science 29(4):530-553.

, and . 1982. Interrelationships of and within Breviraja based on anatomical structures

(Pisces: Rajoidei).—Bulletin of Marine Science, in press.

Stehmann, M. 1970. Vergleichend morphologische und anatomische Untersuchungen zur Neuord-
nung der Systematik der nordostatlantischen Rajidae.—Archiv fur Fischeriewissenschaft
21:73-164.

Whitley, G. P. 1939. Taxonomic notes on sharks and rays.—Australian Zoologist 9:227—262, 2 plates.

. 1940. The fishes of Australia. Part I. The sharks, rays, devil fishes, and other primitive fishes

of Australia and New Zealand. Royal Zoological Society, New South Wales, Sydney. 279 pp.

Department of Wildlife and Fisheries Sciences, Texas A&M University, Col-
lege Station, Texas 77843

PROC. BIOL. SOC. WASH.
95(1), 1982, pp. 13-26

A NEW SPECIES OF GYMNURE, PODOGYMNURA,
(MAMMALIA: ERINACEIDAE) FROM
DINAGAT ISLAND, PHILIPPINES

Lawrence R. Heaney and Gary S. Morgan

Abstract.—A new species, Podogymnura aureospinula, of Philippine gymnure
is described from Dinagat, a small island off the northeast coast of Mindanao in
the southern Philippines. This species is distinguished from other members of the
subfamily by the possession of spiny pelage, inflation of the frontal region, and
presence of a distinct cusp at the base of the talonid notch. This new species is
second in size among the living Echinosoricinae only to Echinosorex gymnurus.
The relationship of Podogymnura to the other extant genera of echinosoricines
is discussed. Based on these comparisons, Podogymnura and Echinosorex are
shown to share a number of cranial and dental characters and are considered to
be more closely related to one another than either is to Hylomys or Neotetracus.

In 1972 and 1975, Dioscoro S. Rabor and a field party from Mindanao State
_University collected mammals and birds on Dinagat and Siargao islands, which
are located off the northeast coast of Mindanao in the southern Philippines. The
itinerary, habitat descriptions, and a report on the birds from the 1972 trip may
be found in duPont and Rabor (1973), and a report on the mammals in Heaney
and Rabor (1982). Specimens from this collection are housed in the Delaware
Museum of Natural History, University of the Philippines at Los Banos, and the
U.S. National Museum of Natural History. Among the mammals collected on
Dinagat are four specimens of a unique member of the family Erinaceidae. In this
paper, we describe these specimens as a new species and discuss the relationships
of the Philippine gymnures, Podogymnura, within the erinaceid subfamily Echi-
nosoricinae.

Methods

External measurements were taken from collector’s labels. Cranial measure-
ments (Table 1) were taken by Heaney with dial calipers graduated to 0.1 mm.
Dental measurements (Table 2) were taken by Morgan using an Anderson cra-
niometer attached to a Bausch and Lomb binocular microscope (Anderson 1968).
All cranial measurements are as defined in DeBlase and Martin (1974) except the
following: rostral length, from midline at anterior tip of nasals to orbital margin
of infraorbital canal; rostral breadth, taken at labial edge of premaxillae just
posterior to I?; post-palatal depth, depth of cranium measured at the point just
posterior of palate to point above at 90° to occlusal plane of molars; I' to M°,
maximum labial length from anterior edge of I' alveolus to posterior edge of M?®
at alveolus; M? to M?, greatest width of palate taken at labial margin of alveoli;
palatal width at M?, alveolar distance between lingual margins; height of coronoid,
maximum vertical height from ventral edge of mandible to tip of coronoid process;
depth of mandible, vertical height from ventral edge of mandible to alveolar

14 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

surface between M, and M,; thickness of mandible, distance from lingual to labial
edge of mandible taken between M, and M,. All dental measurements in Table
1 are dimensions of the teeth, not their alveoli, and represent the maximum length
and/or width of a specified tooth or of a series of teeth. Dental nomenclature
follows Szalay (1969:202). Specimens from the following museums were used in
this study (standard acronym follows in parentheses): Delaware Museum of Nat-
ural History (DMNH), Field Museum of Natural History (FMNH), University of
the Philippines at Los Banos, Museum of Natural History (UPLB), and the U.S.
National Museum of Natural History (USNM).

Podogymnura aureospinula, new species

Holotype.—DMNH 4386, adult female, skin and skull. Obtained 23 April 1972
by Dioscoro S. Rabor (original number 259) at Plaridel, Albor Municipality,
Dinagat Island, Surigao del Norte Province, Republic of the Philippines. Skin well
prepared and in good condition. Skull complete except for zygomatic arches,
both of which are broken (Figs. 1, 2).

Referred specimens.—The holotype and one adult male from Balitbiton, Loreto
Municipality, Dinagat (UPLB 3753) were examined and measured. Two additional
specimens (one male, one female) from Kambinlio, Loreto Municipality, Dinagat,
are in the collection at UPLB, but were not examined.

Diagnosis.—Size large; dorsal pelage short and spinous, golden brown color
overall; temporal, sagittal, and nuchal crests prominent; frontal region conspic-
uously inflated; interorbital region strongly constricted; external pterygoid pro-
cesses large and separated at base from internal pterygoid processes by deep
groove; mandibular rami robust; P? large; P* broad lingually; metaconule prom-
inent on M! and M?; metacone present on M?®; P, with small, but distinct, meta-
conid; distinct cusp at base of talonid notch between metaconid and entoconid
on M,—M,.

Description.—Size (Tables 1 and 2) large for extant members of subfamily.
Dorsal pelage composed of three types of hairs: slate-gray underfur ca. 5 mm in
length; stiff, bristly, or spiny hairs, black at base and remainder golden yellow,
many with black tips, ca. 15 mm in length; black, spiny hairs, ca. 12 mm in
length. Golden spines twice as abundant as black hairs in middle of back. Black
spines densest at mid-dorsum, decreasing in abundance laterally, disappearing on
sides. Black-tipped golden spines especially common at mid-dorsum, also dis-
appearing on sides. Only golden spines present on sides. Overall color of dorsum
golden-brown, with black spines and black-tipped golden spines adding a black
speckling. On holotype, golden color distinctly metallic when viewed at proper
angle. Other specimens somewhat faded, not as metallic. Ventral pelage lacks
spines, grades evenly from dorsal color to brownish-gray over most of venter;
throat darker on some specimens. Ventral hairs of two types: soft, gray underfur
ca. 5 mm in length, and slightly coarser guard hairs ca. 9 mm in length, gray at
base and tipped with light brown. Pelage of rostrum and around eyes short, dense,
and spiny. Upper and lower lips clothed in very short, moderately dense, white
or light brown fur. Vibrissae dark at base, very light for most of length, up to 55
mm. Rhinarium long, naked, and distinctly bilobed, with nostrils opening later-
ally. Ears relatively large, appearing naked, but with sparse covering of extremely

VOLUME 95, NUMBER 1 15

Fig. 1. From top to bottom—dorsal, ventral, and left lateral views of cranium and lateral view of
mandible of Podogymnura truei truei (FMNH 61453) from Mt. Apo, Davao Prov., Mindanao (1-4)
and Podogymnura aureospinula (DMNH 4386), holotype, from Plaridel, Dinagat, Surigao del Norte
Prov. (5-8). Actual size.

Short, white hairs. Fore and hind feet with moderate covering of short, white or
light brown hairs dorsally, nearly naked ventrally. Hind legs appear almost naked
distal to knee joint. Dorsal base of tail with area sparsely furred, nearly naked,
15 mm in diameter. Tail with sparse covering of short hairs. Two pairs of mam-
mae, one pair pectoral, one pair inguinal.

Skull large and robust. Temporal crests converge at or slightly anterior to

16 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Stereophotographs of upper (1, 1’) and lower (2, 2') dentitions of Podogymnura aureo-
spinula (DMNH 4836), holotype. x3.

interorbital constriction to form sagittal crest. Sagittal crest prominent, 0.5 to 1.0
mm in height anteriorly to 2.5 mm in height posteriorly where it meets nuchal
crest. Nuchal crest well developed. Frontal region conspicuously inflated. Fron-
tals expanded laterally into orbital fossa. Rostrum relatively broad posteriorly,
resulting from inflation of posterodorsal portion of maxilla. Inflation causing con-
vexity in dorsal profile of skull, beginning at anterior edge of frontals, approxi-
mately dorsal to P?, and extending posteriorly to interorbital constriction. Highest
point on skull dorsal to orbits. Interorbital region constricted. Braincase relatively
long and not noticeably inflated. Paroccipital and mastoid processes prominent.
Mastoid exposure on ventrolateral corner of skull gently concave and composed
of thick bone. Post-tympanic process of squamosal large, broad posteriorly, and
with well developed epitympanic sinus. Periotic not inflated. Periotic component
of bulla reduced, having a distinct, rounded emargination in ventromedial edge.
Basisphenoid component of bulla more vertically oriented, projecting ventral to

VOLUME 95, NUMBER 1 167

level of occipital condyles. Tympanic cavity broadly open ventrally. External
pterygoid processes large, triangular-shaped, and separated at base from internal
pterygoid processes by deep groove. Mandibular rami relatively thick.

All incisors single-rooted, I’ enlarged. I? and I? much smaller, nearly identical
in size. Canine double-rooted, long, and flared laterally. P? single-rooted, small
compared to other premolars. P? three-rooted, some specimens with slight lingual
expansion and small, but distinct, hypocone. P* broad lingually, with hypocone
and protocone parallel to palatal midline. Metaconule prominent on M? and M?,
protoconule absent. M? relatively large, with small, but distinct metacone. I, and
I, approximately same size, spatulate, and procumbent. I, considerably smaller.
Canine large and vertical, highest tooth in lower tooth row. P, and P; compara-
tively robust, P, one-half the size of P,. P, large, talonid basin moderately to well
developed, and distinct metaconid present. Lower molars with small cusp at base
of talonid notch between entoconid and metaconid, sometimes absent on Ms.
Postcristid on M, and M, slopes lingually at hypoconulid to meet moderately to
strongly developed postcingulid.

Etymology.—L. aureus, golden; L. spinula, diminutive of thorn. The specific
name refers to the golden spines which characterize the dorsal pelage of this
species. We suggest ‘“‘golden-spined gymnure’”’ as an English name.

Comparisons.—Podogymnura aureospinula is more closely related to Podo-
gymnura truei than to any other species in the Echinosoricinae as judged by the
combination of the following characters: long rostrum, absence of postorbital
processes, constricted interorbital region, extreme anterior placement of upper
molariform teeth relative to orbit and infraorbital foramen, relatively small I’, I?
and I? equal in size, large laterally flaring upper canines, loss of P!, comparatively
large P?, and M! and M? square in outline. On the other hand, P. aureospinula
possesses at least three derived characters that are unique among the Echino-
soricinae: spiny dorsal pelage, conspicuously inflated frontal region, and presence
of a cusp at base of talonid notch on lower molars. These characters might justify
generic distinction for P. aureospinula; however, rather than erect a monotypic
genus, we choose to place this new species in the genus Podogymunura to indicate
the presumed monophyletic nature of the two endemic Philippine erinaceids.

Podogymunura truei is the only echinosoricine which requires detailed compar-
ison with P. aureospinula. The other extant echinosoricine genera are compared
to Podogymnura in more general terms in the Discussion section. Podogymnura
aureospinula and P. truei are quite different in external appearance. Besides its
smaller size, P. truei has a longer, softer pelage, with no indication of spines, or
even coarse hairs. Its underfur is particularly long, about twice as long as that of
P. aureospinula. The dorsal surfaces of the fore and hind feet of P. truei have
longer, darker hairs. The overall color of the pelage is darker in P. truei; dorsally
it varies from reddish to chestnut brown compared to the golden brown color of
the large species, and ventrally it is a medium brown, whereas the venter of P.
aureospinula is gray, with a brownish tinge.

The most obvious difference between the skulls of Podogymnura aureospinula
and P. truei is greater size of the former, its skull being 20% longer than the
largest skull of P. truei measured (Table 1). Perhaps as a result of allometric
changes correlated with increasing skull size, the temporal, sagittal, and nuchal
crests and paroccipital and mastoid processes are more prominent in P. aureo-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

18

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VOLUME 95, NUMBER 1

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20 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Selected dental and mandibular measurements (mm) of Podogymnura species.

Podogymnura Podogymnura Podogymnura Podogymnura

aureospinula truei truei truei truei truei minima
Measurement holotype, UPLB 3753 _ holotype (N = 5) (N = 45)

Length of maxillary tooth row 20.6, 20.4 [52 15.5 + 0.44 15.0 + 0.65

(15.1—16.2) (14.4-15.9)

Length from M! to M? 9.8, 9.6 6.7 ies 0i20 7.0 + 0.33
(6.9-7.4) (6.6~7.5)

Length of P® Me PES) 1.8 19 1050 1.8 + 0.08
(1.8—2.1) (1.7—1.9)

Width of P® ORAS MTT, ell ese = 017. 1.4 + 0.07
(1.2-1.6) (1.3—1.5)

Length of P* 37, 310 D2 2.4 + 0.05 mse (Ibi
(2.3-2.4) (2:3=2.6)

Width of P* 3.2, 3.0 DN 2.4 + 0.10 2.4 + 1.54
(2.3—2.5) (2.2—2.5)

Length of M! 3.8, 3.8 2.6 2.9 + 0.04 2.8 + 0.11
(2.82.9) (2.62.9)

Width of M! alls St! 2.3 Deh = On 2.6 + 0.08
(2.62.9) (2.5-2.7)

Length of M? BWo)5 Bias) 2.4 2°) 20S 2252-016
(2.42.7) (2.3—2.7)

Width of M? 3.7% 33 2.4 DEG, == 10205 Donne lel
(2.6—2.7) (2.5—2.7)

Length of M? hs, 2D led 1.9) = O15 1.8 + 0.09
(1.7—2.1) (1.7—1.9)

Width of M?® DES OES 1.8 2.1 + 0.08 2.0r=,0 eli
(2.0-2.2) (1.9-2.1)

Total length of mandible 41.2, 40.8 B30 e077. 28.8 + 0.58
(29.6—31.5) (28.2-29.6)

Height of coronoid 13)... 113).4! 9.2 9.7 + 0.50 9.3 + 0.59
(9.0-10.4) (8.6—10.0)

Depth of ramus between M, and M, 4.7, 4.8 Zall 3 op = OR2 Bop eas | )e75)
(3.0-3.6) (2.8-3.5)

Breadth of ramus between M, and 2.6, 2.8 1.6 1.8 + 0.05 1.7 + 0.07
M, (1.7—1.8) (1.6+1.8)

Length of mandibular toothrow DNA 207. 15.8 1622) 310354 15.7 + 0.64
(15.7=17 1) (15.1—-16.6)

Length from M, to M, 11.2, 10.5 7.8 8.0 + 0.26 7.8) 10
(7.6-8.2) (7.5-8.3)

Length of P, 3.0, 2.6 1.9 2.0 + 0.10 2.0 + 0.19
(129-771) (1.8-2.3)

Width of P, IRS, les) ok 1.3 + 0.04 12008
(1.2=1.3) (1.1-1.3)

Length of M, 4.3, 4.1 3.0 Byaa | ree (0a 2.9 + 0.09
(39-3) (2.9-3.2)

Width of M, RSE on 5) loi 1.9+ 0.13 1.8 + 0.04
(1.7—2.0) (1.8-1.9)

VOLUME 95, NUMBER 1 21

Table 2.—Continued.

Podogymnura Podogymnura Podogymnura Podogymnura

aureospinula truei truei truei truei truei minima
Measurement holotype, UPLB 3753 holotype (N = 5) (N = 4-5)

Length of M, 4105 Sal! Met 2.8 + 0.13 2.7 + 0.08
(2.62.9) (2.6-2.8)

Width of M, 2.4, 2.4 1.6 18+ 0.11 1.8 + 0.09
(1.7-1.9) (1.71.9)

Length of M, 33.54 De 2.3+0.11 2.3 + 0.15
(22214) (Qal=255)

Width of M, Delle, PAY 1.4 1255-70505) ls) ee LNG

(1.5-1.6) (1LZ=LD)

spinula, approaching the condition seen in the much larger Echinosorex. In most
skulls of P. truei, the weak temporal crests meet near the anterior edge of the
interparietal, forming a short, weak sagittal crest, while in larger skulls of the
Same species, the temporal crests meet somewhat farther forward. The temporal
crests of P. aureospinula converge at or slightly anterior to the interorbital con-
striction to form the strong sagittal crest, which is particularly high posteriorly
where it bisects the interparietal. The nuchal crest of P. aureospinula is more
strongly developed than in P. truei.

Podogymuura truei shows no evidence of the frontal inflation characteristic of
P. aureospinula. The dorsal margin of the skull of P. truei rises in a nearly
straight line to reach a maximum height above the glenoid region, rather than
dorsal to the orbits as in P. aureospinula. The inflation of the frontals and pos-
terodorsal portion of the maxilla of P. aureospinula is not characteristic of any
other modern echinosoricine, although some erinaceines (e.g. Paraechinus) have
moderately inflated frontals. The interorbital region appears to be more constrict-
ed in P. aureospinula, although this feature is certainly enhanced by the inflation
anterior to the constriction. The more prominent development of the sagittal and
nuchal crests gives the braincase of P. aureospinula the appearance of being
longer, whereas the braincase of the smaller species appears shorter, broader,
and more bulbous, particularly in smaller specimens.

The mastoid exposure on the posterolateral corner of the skull is inflated in
Podogymuura truei and the bone is thin and nearly transparent. The mastoids of
P. aureospinula are gently concave and composed of thicker bone. The periotic
of P. truei is also inflated. The periotic is not inflated in P. aureospinula and the
periotic component of the bulla has a small, rounded emargination in its ventro-
medial edge which is lacking in the smaller species. The basisphenoid portion of
the bulla of P. aureospinula is more vertically oriented, projecting ventral to the
occipital condyles, whereas the basisphenoid bulla of P. truei is more horizontal,
tending to enclose the tympanic cavity. Coupling the more vertical orientation of
the basisphenoid bulla with the reduced periotic portion of the bulla, the tympanic
cavity is more open ventrally in P. aureospinula. The post-tympanic process of
the squamosal is relatively larger in P. aureospinula than in any other modern
echinosoricine. Its external pterygoid processes are also relatively large, but these
are smaller in P. truei. Podogymnura aureospinula has a deep groove at the base

22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

of the external pterygoid processes which separates them from the internal pter-
ygoid processes. There is no evidence of this groove in any specimen of P. truei
examined. As with the skull, the mandible of P. aureospinula is larger and more
robust than that of its smaller counterpart. The mandibular rami are particularly
thickened in P. aureospinula.

The upper incisors and canines of Podogymnura aureospinula and P. truei are
similar in morphology. The P? is somewhat longer and broader in P. aureospinula;
it is more conical in P. truei. In most specimens of P. truei, the P® is two-rooted,
thin, and blade-like, with no indication of a protocone or hypocone. Butler (1948)
stated that the P? of Podogymnura had only two roots; however, the larger series
of specimens now available shows this character to be variable. Most specimens
of P. truei have a two-rooted P®, but the few that have a slight lingual expansion
of P® have three roots. The P® of P. aureospinula is three-rooted, and bears a
small hypocone. The P* of P. aureospinula is broader lingually, the hypocone is
larger, and the protocone is higher and more lingually placed. In comparison, the
P* in the smaller species is rather narrow lingually, the hypocone is small, and
the protocone is located closer to the paracone. The M! and M? are similar in the
two species, except for the more prominent metaconule and cingula in P. au-
reospinula. The M? of P. aureospinula is larger and bears a small metacone, the
latter being absent on the M? of P. truei.

As in the upper dentition, the P, and P, are relatively larger and more robust
in P. aureospinula than in P. truei. The talonid basin is reduced on P, in P. truei
and a metaconid is absent or tiny. The P, of P. aureospinula has a moderately
to well developed talonid basin and the metaconid is distinct. The morphology
of the lower molars is similar in the two species. P. aureospinula has a small,
distinct cusp located at the base of the talonid notch between the entoconid and
metaconid on M,, M,, and sometimes M,;. The entoconulid is normally located
slightly anterior to, but close to the entoconid, and therefore this cusp does not
appear to be homologous with an entoconulid. This cusp is absent in P. truei and
seems to be unique among modern echinosoricines.

Ecology.—Virtually nothing is known of the natural history of Podogymnura
aureospinula. The vicinity where the holotype of P. aureospinula was taken was
described by duPont and Rabor (1973:4) as *‘. . . a logged area in rolling country
and low hills where there were still many patches of remnant dipterocarp forests
in the surrounding localities.’’ These forests are dominated by Dipterocarpus,
Shorea, Hopea, Anisoptera, and Pentacme among the Dipterocarpaceae, and
include members of at least eleven other plant families. Undergrowth consists
mainly of rattan and ferns. Other terrestrial mammals taken at the type locality
of P. aureospinula include Urogale everetti, Tarsius syrichta, Cynocephalus vo-
lans, Sundasciurus mindanensis, Exilisciurus surrutilus, Batomys sp., Rattus ev-
eretti, and Rattus rattus (Heaney and Rabor 1981).

Additional specimens examined.—Podogymnura truei truei (2 63, 4 2°).
PHILIPPINES, Mindanao, Davao Province: E slope of Mt. McKinley, 5800 ft.
elev., FMNH 56129, 56172, 56181; N slope of Mt. Apo, Lake Linau, 7800 ft.
elev., FMNH 61435; E slope of Mt. Apo, Baclayan, 5400 ft. elev., FMNH 61453;
Mt. Apo, 6000 ft. elev., USNM 125286 (holotype).

Podogymnura truei minima (2 66,3 2° ¢). PHILIPPINES, Mindanao, Bukid-
non Province: Mt. Katanglad, near Malay Balay, DMNH 5949-5953.

VOLUME 95, NUMBER 1 23

Echinosorex gymnurus albus (1 6, 4 22). INDONESIA, Borneo, Sempang
River: USNM 145581-582, 145584—586.

Hylomys suillus dorsalis (10 63, 1 2). MALAYSIA, Borneo, Sabah, Mt.
Kinabalu, Bundu Tuhan: USNM 292337-339, 292341-342, 292350-354, 292356.

Neotetracus sinensis (1 6, 2 22, 42). CHINA: Yunnan, Ho mu shu Pass,
USNM 241402; Szechuan, Kwan Shien, 3000 ft. elev., USNM 258124-129.

Discussion

The genus Podogymnura and its type species, P. truei, were described by
Mearns (1905) from a single specimen collected on Mount Apo, south-central
Mindanao, in the Philippines. Until the late 1940’s, this specimen, an adult female
consisting of a complete body in alcohol and a skull lacking the zygomatic arches
and braincase, was the only known specimen of the genus. Mearns provided a
detailed description of the external characters of P. truei but he did not illustrate
the cranium or mandible and his description of them was brief. He described the
dentition of P. truei, comparing it with that of Hylomys, but his descriptions and
comparisons are so general that they are of little use. Lyon (1909) noted that
although Podogymnura and Hylomys appeared to be closely related with respect
to size and external characters, they were distinct dentally. Cabrera (1925) pro-
vided a key to the living echinosoricine genera in which he distinguished Podo-
gymnura by the combination of loss of P}, P*® larger than P?, and larger upper
canines. Butler (1948) regarded Podogymnura as intermediate between Echino-
sorex and the smaller Hylomys and Neotetracus, although he pointed out that in
the enlargement of the canines, relatively large P®, length of the rostrum, and
position of P* and M' relative to the orbit and infraorbital foramen, Podogymnura
is similar to Echinosorex.

Sanborn (1952) reported on 64 specimens of Podogymunura truei collected on
Mount Apo and Mount McKinley on Mindanao. He compared Podogymuura to
the other genera of modern echinosoricines, reaching the same general conclusion
as did Butler. Sanborn (1953) described a new subspecies of Podogymnura truei,
P. t. minima, from four specimens collected on Mount Katanglad, Bukidnon
Province, north-central Mindanao. His diagnosis was based exclusively on the
smaller size of P. t. minima. Detailed cranial and dental measurements of P. f.
truei (including the type) and topotypic specimens of P. t. minima (Tables | and
2) indicate that, on the whole, the available specimens from Mount Katanglad are
slightly smaller than specimens from Mount Apo and Mount McKinley, but there
is broad overlap in size between them, especially in dental measurements. In
fact, the teeth of the type specimen of P. t. truei from Mount Apo are actually
smaller in many dental measurements than the teeth of the topotypic specimens
of P. t. minima. Sanborn specifically noted that there was no difference in color
between his series of P. t. minima and the nominate form. The DMNH specimens
of P. t. minima, however, are lighter in color than topotypic specimens of P. f.
truei, being slightly more reddish brown and having the tail uniformly lighter.
Besides the references cited above, Hollister (1913), Taylor (1934), and Alcasid
(1970) mentioned Podogymnura, but provided no new information.

Comparison of Podogymnura with other living echinosoricines.—Butler (1948)
made exhaustive comparisons of three of the five extant genera of echinosori-

24 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

cines: Hylomys, Neotetracus, and Echinosorex. His comparisons of Podogym-
nura were general, as they were based solely on published descriptions and fig-
ures of the type of P. truei, the only specimen of the genus known at that time.
The fifth living genus of echinosoricine, Neohylomys, was not described until
1959. Since 1948 many specimens of Podogymnura have been collected, but the
genus has not been adequately compared with the other members of its subfamily.
Our description of a new species of Podogymnura, which in some characters
appears to bridge the morphological gap between Podogymnura and Echinosorex,
calls for fuller comparisons and a reassessment of the phylogenetic relationships
of the Philippine gymnures.

As Butler (1948) and others have noted, Hylomys and Neotetracus are closely
related forms, although in our opinion they do represent distinct genera, contrary
to Van Valen (1967). The status of Neohylomys is uncertain; the only specimens
are in China, and unavailable for study. Many of the characters unique to Hy-
lomys and Neotetracus, such as the reduced antemolar dentition, the palatal
perforations, the presence of posterior processes of the maxillae and anterior
processes of the parietals which extend across the frontals and meet or nearly
meet dorsal to the orbits, the prominent flange on the anterior edge of the orbit,
and the strongly concave anterior portion of the zygomatic arch for attachment
of the lateral snout muscles, are derived characters which demonstrate a close
phylogenetic relationship between the two genera. Conversely, many of the char-
acters that are shared by Podogymnura and Echinosorex, but are not present in
the other two genera, appear to be primitive for the Echinosoricinae. Of these
characters, the longer rostrum, better developed maxillary dentition anterior to
P*, widely separated maxillae and parietals dorsal to the orbits, and the nonper-
forated palate are certainly primitive. In the remaining characters shared by Po-
dogymnura and Echinosorex, it is difficult to determine the primitive character
state for the Echinosoricinae because no appropriate outgroup exists. Butler
(1948) felt that the two distinct groups of living echinosoricines represented two
separate specialized offshoots from the primitive condition. Suffice it to say that
Podogymnura and Echinosorex share so many cranial and dental features that
they undoubtedly represent closely related genera.

Although they share many cranial characters, Podogymnura and Echinosorex
are extremely different externally. Echinosorex is a larger animal with a tail equal
to about 60% of the length of head and bedy. Podogymnura is medium-sized (P.
aureospinula) or small (P. truei) and has a tail equal to about 35% of the length
of head and body. Podogymuura is a chestnut brown (P. truei) to golden brown
(P. aureospinula) color and has relatively short pelage. Echinosorex is either pure
white (E. gymnurus albus) or predominantly black with white markings on the
neck and face (all other forms of the species) and has long, coarse guard hairs on
the dorsum. Based on its external appearance, especially its small size and short
tail, Podogymnura truei more closely resembles Hylomys and Neotetracus than
Echinosorex.

Additional cranial characters shared by Podogymnura and Echinosorex not
listed above include: well developed canines and P®, small supraorbital crests,
presence of two longitudinal grooves in maxillary component of hard palate which
extend from incisive foramina to small foramina located medial to P? or P®, pres-

VOLUME 95, NUMBER 1 25

ence of a ventral process on maxillary portion of zygomatic arch, and more
anterior placement of upper cheek teeth relative to orbit and infraorbital foramen.
The tympanic bone is slender and not firmly attached to the bulla in Podogymnura
and Echinosorex, whereas in Hylomys and Podogymnura the tympanic is broader
and is firmly attached to the edge of the bulla, contributing to its formation and
restricting the size of the opening of the tympanic cavity. Although Butler (1948)
stated that Echinosorex lacked the anterior process of the tympanic, the large
series of echinosoricines at our disposal reveals that both Echinosorex and Po-
dogymnura definitely have an anterior process, but it is smaller than that of
Hylomys and Neotetracus.

In addition to the differences in external morphology cited above, there are a
number of cranial and dental characters that are diagnostic of Podogymnura and
distinguish it readily from Echinosorex. These include: less prominent temporal,
sagittal, and nuchal crests and mastoid and paroccipital processes; more concave
maxillary portion of zygomatic arch; broader, more inflated braincase; more an-
terior placement of upper molars relative to orbit and infraorbital foramen (M? is
anterior to front edge of orbit and M! is below infraorbital foramen); smaller I;
I? and I*® equal in size; canines relatively larger and flared laterally; loss of P};
M? square in outline; absence of protoconule on M!; smaller metaconule on M!
and M?; reduced posterolingual apex of M?.

Podogymnura aureospinula is intermediate between Echinosorex and P. truei
in many characters, although some of the characters on which this observation
is based are probably related to its intermediate size. The more prominent tem-
poral, sagittal, and nuchal crests and mastoid and paroccipital processes of P.
aureospinula relative to P. truei are almost certainly correlated with its greater
size. These features of the bony crests and processes are best developed in Echi-
nosorex, the largest genus among living echinosoricines, and are likely to be
allometric changes associated with increasing skull size. Other intermediate char-
acters of P. aureospinula which are more difficult to ascribe to allometry are the
larger P?, more prominent metaconule on M! and M?, the presence of a metacone
on M?, and the deep groove separating the bases of the external and internal
pterygoid processes. Whether these are derived characters indicating a closer
relationship between Echinosorex and P.. aureospinula or whether these represent
primitive characters shared by Echinosorex and P. aureospinula and lost by P.
truei cannot be determined from available data. Regardless of the similarities
between P. aureospinula and Echinosorex, the two species of Podogymnura are
certainly more closely related to one another than either is to Echinosorex and
have probably been isolated in the Philippines for a considerable period of time.

Acknowledgments

We wish to thank M. D. Carleton, P. Myers, and R. W. Thorington for valuable
comments on the manuscript, and G. Lake and S. F. Campbell for assistance in
preparation of the manuscript and tables. Photographic assistance was provided
by D. Bay and V. Krantz. We thank P. Freeman, D. Niles, D. S. Rabor, R. W.
Thorington, and R. M. Timm for permission to examine specimens under their
care.

26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Literature Cited

Alcasid, G. L. 1970. Checklist of Philippine mammals.—National Museum, Pee of Education
(Manila). Museum Publication No. 5:1-51.

Anderson, S. 1968. A new craniometer and suggestions for craniometry.—Journal of Mammalogy
49:221-228.

Butler, P. M. 1948. On the evolution of the skull and teeth in the Erinaceidae, with special reference
to fossil material in the British Museum.—Proceedings of the Zoological Society of London
118:446—S00.

Cabrera, A. 1925. Genera Mammalium—Insectivora, Galeopithecia——Museo Nacional Ciencia
Natural, Madrid, 232 pp.

DeBlase, A. F., and R. E. Martin. 1974. A manual of mammalogy. Wm. C. Brown Co., Dubuque,
Iowa, 329 pp.

duPont, J. E., and D. S. Rabor. 1973. Birds of Dinagat and Siargao, Philippines, an expedition
report.—Nemouria (Occasional Papers of the Delaware Museum of Natural History) 10:1-111.

Heaney, L. R., and D. S. Rabor. 1982. An annotated checklist of the mammals of Dinagat and
Siargao islands, Philippines.—(Occasional Papers of the Museum of Zoology, University of
Michigan) (In press.)

Hollister, N. 1913. A review of the Philippine land mammals in the United States National Museum.—
Proceedings of the United States National Museum 46:299-341.

Lyon, M. W., Jr. 1909. Remarks on the insectivores of the genus Gymnura.—Proceedings of the
United States National Museum 36:449_456.

Mearns, E. A. 1905. Descriptions of new genera and species of mammals from the Philippine Is-
lands.—Proceedings of the United States National Museum 28:425—460.

Sanborn, C. C. 1952. Philippine zoological expedition 1946-47: Mammalis.—Fieldiana: Zoology

33(2):89-158.

. 1953. Mammals from Mindanao, Philippine Islands collected by the Danish Philippine Ex-

pedition 1951—1952.—Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening

115:283-288.

Szalay, F. S. 1969. Mixodectidae, Microsyopidae, and the insectivore-primate transition.—Bulletin
of the American Museum of Natural History 140:193-330.

Taylor, E. H. 1934. Philippine land mammals.—Monographs of the Bureau of Science, Manila
30: 1-548.

Van Valen, L. 1967. New Paleocene insectivores and insectivore classification.—Bulletin of the
American Museum of Natural History 135:217-284.

(LRH) Museum of Zoology and Division of Biology, University of Michigan,
Ann Arbor, Michigan 48109; (GSM) Division of Mammals, U.S. National Mu-
seum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

Present address of GSM: Florida State Museum, University of Florida, Gaines-
ville, Florida 32611.

PROC. BIOL. SOC. WASH.
95(1), 1982, pp. 27-47

CHECKLIST OF THE FISHES OF THE CENTRAL AND
NORTHERN APPALACHIAN MOUNTAINS

Jay R. Stauffer, Jr., Brooks M. Burr, Charles H. Hocutt, and
Robert E. Jenkins

Abstract.—A table lists 398 forms and 5 intergrade populations in 28 families
in an area on the Atlantic slope from the Susquehanna River south to the Peedee
River, including Ohio River basin drainages from the Monongahela River in Penn-
sylvania to the Tennessee River in Alabama and Tennessee.

The central Appalachians harbor a diverse fish fauna that includes numerous
endemics. Jenkins, Lachner, and Schwartz (1972), as part of a zoogeographic
analysis of this ichthyofauna, provided a table that lists the fishes of the central
Appalachians by river drainage and general habitat. This table has been extremely
valuable to ichthyologists, fisheries scientists, and environmental consultants
throughout the past decade.

Numerous studies have substantially increased our knowledge of fish distri-
bution throughout the central Appalachians (Hambrick et al. 1973, Hocutt and
Hambrick 1973, Hocutt et al. 1973, Stauffer et al. 1975, Stauffer et al. 1976,
Hocutt et al. 1978, Stauffer et al. 1978, Hendricks et al. 1979, Hocutt et al. 1979,
Lee et al. 1980) and indicated that the original table should be updated. Moreover,
it was thought that the addition of the Susquehanna, Licking, Green, and Ken-
tucky rivers would enhance the usefulness of the faunal list.

The list (Table 1) includes 398 forms and 5 intergrade populations in 28 families.
It covers an area on the Atlantic Slope from the Susquehanna River (New York
and Pennsylvania) south to the Peedee River (North Carolina and South Caroli-
na). Ohio River basin drainages that are included extend from the Monongahela
River in Pennsylvania south to the Tennessee River in Alabama and Tennessee.

It should be noted that the list is conservative. If a question exists as to the
current or historic presence of a species, it is not included. No attempt is made
to distinguish species that were historically present in the drainage from those
that currently occur. Trinomials are used only when the distribution of subspecies
could be accurately determined.

The authors appreciate the encouragement of Dr. E. Lachner, who recognized
the need for a revision of the original faunal list.

28 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Fishes of the central and northern Appalachian drainages and their general habitat. KX =
category. most frequently inhabited. Drainage occurrence: E = endemic, N = native, NP = probably
present—native, NI = regarded as native but possibly introduced, I = introduced, IP = probably or
possibly introduced, Ma = euryhaline or diadromous. Native extralimital distribution: So = south on
Atlantic slope, No = north on Atlantic slope, O = predominantly Ohio basin form, M = lower and/or
central Mississippi basin, G = Gulf of Mexico slope.

PETROMYZONTIDAE

Ichthyomyzon bdellium
Ichthyomyzon castaneus
Ichthyomyzon fossor
Ichthyomyzon gagei
Ichthyomyzon greeleyi
Ichthyomyzon unicuspis
Lampetra aepyptera
Lampetra appendix
Petromyzon marinus

ACIPENSERIDAE

Acipenser brevirostrum
Acipenser fulvescens
Acipenser oxyrhynchus
Scaphirhynchus platorynchus

POLYODONTIDAE
Polyodon spathula
LEPISOSTEIDAE

Lepisosteus spatula
Lepisosteus oculatus
Lepisosteus osseus
Lepisosteus platostomus

AMIIDAE

Amia calva

ANGUILLIDAE

Anguilla rostrata

CLUPEIDAE

Alosa aestivalis

Alosa alabamae

Alosa chrysochloris
Alosa mediocris

Alosa pseudoharengus
Alosa sapidissima
Dorosoma cepedianum
Dorosoma petenense

HIODONTIDAE

Hiodon alosoides
Hiodon tergisus

SALMONIDAE

Coregonus artedii
Coregonus clupeaformis

Lowland

x x KK

x mK x

x KK KKK OM

x xX

Upland

mK KK KK

x x

Habitat

Montane
Big River

~*~ x

xxx KKK OK * x mK mK va x Km

x x

Stream

KK mK KK mK OK OM OM

Creek

x xX

Peedee

Cape Fear

Ma

Ma

Neuse

Drainage occurrence

Atlantic Slope

Tar

Roanoke

NN <Ne

Ma

Ma

Ma

Ma

Ma

Ma
Ma
Ma
Ma

Ma

Ma

James

York

Rappahannock

Potomac

Susquehanna

Ne NENG ENGIN
N N N N N
Ma Ma Ma Ma Ma Ma

Ma

Ma

Ma

Ma

Ma

Ma

Ma

Ma

NP

Ma

Ma

Ma

Ma

Ma

Ma

Ma

Ma

Ma

Ma

Ma

Ma

IP

VOLUME 95, NUMBER 1

Table 1.—Continued.

29

PETROMYZONTIDAE

Ichthyomyzon bdellium
Ichthyomyzon castaneus
Ichthyomyzon fossor
Ichthyomyzon gagei
Ichthyomyzon greeleyi
Ichthyomyzon unicuspis
Lampetra aepyptera
Lampetra appendix
Petromyzon marinus

ACIPENSERIDAE

Acipenser brevirostrum
Acipenser fulvescens
Acipenser oxyrhynchus
Scaphirhynchus platorynchus

POLYODONTIDAE
Polyodon spathula

LEPISOSTEIDAE

Lepisosteus spatula
Lepisosteus oculatus
Lepisosteus osseus
Lepisosteus platostomus

AMIIDAE

Amia calva

ANGUILLIDAE

Anguilla rostrata

CLUPEIDAE

Alosa aestivalis

Alosa alabamae

Alosa chrysochloris
Alosa mediocris

Alosa pseudoharengus
Alosa sapidissima
Dorosoma cepedianum
Dorosoma petenense

HIODONTIDAE

Hiodon alosoides
Hiodon tergisus

SALMONIDAE

Coregonus artedii
Coregonus clupeaformis

(or)
a OS
Dayna
ey
o.6U6U
= 5
NP N
N
N N
N
N WN
Ma Ma
Ma Ma
Ma
N
N

Kanawha: below falls

Ma

Ma

Kanawha: above falls

Drainage occurrence

Ohio Basin
&
6 6s
> ep 3
Ory ye iS
NP NP N
N NP
NP NP
ING NN
N NP
N
N N
Ma Ma Ma
N
Ma Ma
Ma Ma
Ma Ma Ma
N NP
N

Kentucky

Lik LZ,

NP

Green

Z

Z ZZ Z

NP

Ma
Ma

Ma

Ma

Cumberland: below falls

i 7

EG

Cumberland: above falls

Ze Tennessee

Ly Li Law Lie,

Native
extralimital
distribution

Atlantic Slope

NoSo

NoSo

NoSo

NoSo

Elsewhere

ie)

Hae 2

GM

GM

GM

GM

30 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Drainage occurrence

Atlantic Slope

x
tab}
ion
=.
p
+

Lowland
Upland
Montane

Big River
Stream
Creek
Peedee

Cape Fear
Neuse
Roanoke
James

York
Rappahannock
Potomac
Susquehanna

Tar

Oncorhynchus kisutch

Oncorhynchus nerka

Salmo gairdneri I
Salmo trutta I I I I
Salvelinus fontinalis XxX DS DP N N NI

Salvelinus namaycush
OSMERIDAE

Osmerus mordax NI

UMBRIDAE

Umbra limi X Nea OX
Umbra pygmaea 4 XX“ N-N N NUN ON SON NNN

ESOCIDAE
Esox a. americanus xX xX xX N N N -“N N 2INSIN SN NSN
Esox a. vermiculatus > XX
Esox lucius

I

Esox masquinongy Xe XS WX’ I I I
Esox niger xX XxX xX XN N N NeoiNeoN aN eNaaNeeN
I

Esox reicherti

p=
—
—
Za
a” Ail en il onl oon lo

a

—

CY PRINIDAE

Campostoma a. anomalum x OX xX XX IP N N N N
Campostoma oligolepis XE 20 OX Xx

Carassius auratus I I I I
Clinostomus elongatus NI
Clinostomus f. funduloides
Clinostomus f. estor
Clinostomus f. subsp. Xx
Couesius plumbeus
Ctenopharyngodon idella xX xX

Cyprinus carpio rT, 2 ~habe a oe eerie ieee
Ericymba buccata
Exoglossum laurae
Exoglossum maxillingua
Hemitremia flammea
Hybognathus hayi
Hybognathus n. nuchalis
Hybognathus n. regius
Hybopsis aestivalis hyostoma
Hybopsis amblops

Hybopsis cahni

Hybopsis d. dissimilis
Hybopsis hypsinotus
Hybopsis i. insignis
Hybopsis i. eristigma
Hybopsis labrosa

Hybopsis monacha

Hybopsis storeriana XxX
Hybopsis x-punctata

xx x
x KKK

IP

*
*
*

x KK xX
~

x x x

~*~

Z,

Z,

iL,

xx x
x KKK
KKK KKK KKK KK

mK KKK KKK KKK

*

VOLUME 95, NUMBER 1 31

Table 1.—Continued.

Drainage occurrence

Ohio Basin
a I 5 = Native
a8 S$ Gistibution
os 2 S 3) 5
8 Fs oO = . a
EES at accrnrian aie se BS SRG 0 i omar
3 a x 3 5 5 > a 3 o n ©
he ae ag iN pe emg ae 2) a
Seca Se Se Se en ci, Vee, Rok ree Sal ee a 5
So a a Be o o 2 =| =| 5 cs 2)
Se pee Sa Oe eh A Omer Oe < cal
Oncorhynchus kisutch
Oncorhynchus nerka I I
Salmo gairdneri I I I I I I i I I I I
Salmo trutta I I I I I I I I
Salvelinus fontinalis NG OEP NN IP I N NoSo
Salvelinus namaycush No
OSMERIDAE
Osmerus mordax I
UMBRIDAE
Umbra limi N M
Umbra pygmaea NoSo G
ESOCIDAE
_Esox a. americanus I NoSo G
Esox a. vermiculatus New INE EN2 Ne ON N GM
Esox lucius I I I I M
Esox masquinongy No INN Y UIP ye LNG INE ON IN N
Esox niger I I I N N NoSo GM
Esox reicherti
CY PRINIDAE
Campostoma a. anomalum INS dN IND INL TUNG) CINE UING aN N NoSo O
Campostoma oligolepis N, NE VNOOAIN GM
Carassius auratus I I I I I I I I
Clinostomus elongatus N N No OM
Clinostomus f. funduloides N N N N NI So O
Clinostomus f. estor N N
Clinostomus f. subsp. N So
Couesius plumbeus No
Ctenopharyngodon idella
Cyprinus carpio I I I I I I I I I I I I
Ericymba buccata INI SENG Ne INE ONIN NO SING ING DON ONT IN efor GME
Exoglossum laurae NI N O
Exoglossum maxillingua NI No
Hemitremia flammea N N G
Hybognathus hayi N GM
Hybognathus n. nuchalis N ING Ni N GM
Hybognathus n. regius NoSo
Hybopsis aestivalis hyostoma N Ne ONe IN! AN. EN IN N GM
Hybopsis amblops IN NN NN Nie Ni SIN N M
Hybopsis cahni E
Hybopsis d. dissimilis No Ne) NON N N N N N O
Hybopsis hypsinotus So
Hybopsis i. insignis N N
Hybopsis i. eristigma E
Hybopsis labrosa So
Hybopsis monacha E
Hybopsis storeriana N N N NON) ANE Ne EN N GM

Hybopsis x-punctata N M

32 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Drainage occurrence

Atlantic Slope

Habitat

Lowland
Upland
Montane

Big River
Stream
Creek
Peedee

Cape Fear
Neuse

Tar

Roanoke
James

York
Rappahannock
Potomac
Susquehanna

x
*
Z,
Z,

Hybopsis sp. cf. zanema
Leuciscus idus I
Nocomis biguttatus

Nocomis effusus

Nocomis |. leptocephalus

Nocomis 1. bellicus

Nocomis |. interocularis

Nocomis micropogon

Nocomis platyrhynchus

Nocomis raneyi

Notemigonus crysoleucas XxX
Notropis albeolus

Notropis alborus

Notropis altipinnis 4

Notropis amnis xX XxX
Notropis amoenus xX

Notropis analostanus

Notropis ardens

Notropis ariommus

Notropis atherinoides XxX
Notropis baileyi

Notropis bellus

Notropis bifrenatus XxX
Notropis blennius xX X
Notropis boops

Notropis buchanani xX
Notropis camurus x
Notropis cerasinus

Notropis chalybaeus Xx
Notropis chiliticus

Notropis chrysocephalus

Notropis coccogenis

Notropis cornutus

Notropis c. cummingsae xX
Notropis d. dorsalis

Notropis emiliae XxX
Notropis fumeus XxX
Notropis galacturus

Notropis heterodon

Notropis heterolepis

Notropis hudsonius 4
Notropis leuciodus

Notropis lirus

Notropis lutipinnis

Notropis lutrensis XxX
Notropis maculatus x
Notropis mekistocholas

Notropis niveus

Notropis petersoni xX

IP. 7

x KK mK XK
Z
Z
Z
Z
Z
Z
Z

x KX
x x XK

NI

~ KKK KKK KK KK
KKK KK mK KK
Ze
Z,
Z
Z
Z,

xx KK
ZZ,

YL, by Fy TA
Zi ZL LL,

NI

<> > > > >
mM td tO
s >< ><
Z
Zz
A WR We

x KK x
x
x KK XK

IP IP

xx Kx
x Kx
xx KK
xx KKK
ZA
Zz
LZ,
eZ
ZZZ
ZZ

xe KKK KKK CK
*
xx KK OK
FLT 5p TS

x x xX

x x
*
KKK mK KK Km KK KK
~

Pi, 4
74, ZA Nos) ZZ
ZA
Z

VOLUME 95, NUMBER 1 33

Table 1.—Continued.

Drainage occurrence

Ohio Basin
= = : s Be tee
SS oa a > distribution
Es ee Bie =
Ss SS 0) a) 53 a
OReery ne e ee eu ey, a ic eae
S a a a 5 5 > Ss SI 7) 2
ee i Wl ee ete FP SP Bee ine
a x) s x 3 n as} £ © ro as) < S 3
SS se om a SRO, Gh BE ee
Er eee. Nae, OMe i OMA a hy ms | ie < ea)
Hybopsis sp. cf. zanema
Leuciscus idus
Nocomis biguttatus NI M
Nocomis effusus N N N
Nocomis |. leptocephalus N So
Nocomis 1. bellicus N G
Nocomis |. interocularis 2? SO
Nocomis micropogon IN WEN? ON NP NEN UN Ni Ne SNiaeNo O
Nocomis platyrhynchus E
Nocomis raneyi
Notemigonus crysoleucas N Nee NN > N) ¥elyeN) SNoSomGM
Notropis albeolus N
Notropis alborus So
Notropis altipinnis So
‘Notropis amnis N N N GM
Notropis amoenus No
Notropis analostanus No
Notropis ardens N NE INE ON) IN UN oNiN O
Notropis ariommus N N N IND INP INT N O
Notropis atherinoides N N NN NG Ny ON ON EN NE NogGivi
Notropis baileyi N G
Notropis bellus N G
Notropis bifrenatus No
Notropis blennius N N NN ENON UN @N N M
Notropis boops NN NN N M
Notropis buchanani N N NP NG SNew No IN N GM
Notropis camurus N
Notropis cerasinus N
Notropis chalybaeus NoSo GM
Notropis chiliticus IP
Notropis chrysocephalus ING) NING LIBS ONO Ne ON UN ON Ae aNaeN GM
Notropis coccogenis IP N So
Notropis cornutus N N No M
Notropis c. cummingsae So G
Notropis d. dorsalis N
Notropis emiliae N N N So GM
Notropis fumeus N N N GM
Notropis galacturus NI N N IP ON M
Notropis heterodon No
Notropis heterolepis N No M
Notropis hudsonius N IP NoSo M
Notropis leuciodus IP N N N So O
Notropis lirus N G
Notropis lutipinnis IP So
Notropis lutrensis
Notropis maculatus So GM
Notropis mekistocholas
Notropis niveus So
Notropis petersoni So G

SSS

34 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Drainage occurrence

Atlantic Slope

a0)
p
g,
ot
feb}
>

Lowland
Upland
Montane

Big River
Stream
Creek
Peedee

Cape Fear
Neuse

Tar

Roanoke
James

York
Rappahannock
Potomac
Susquehanna

x

Notropis photogenis
Notropis procne procne
Notropis p. longiceps
Notropis pyrrhomelas
Notropis rubellus
Notropis rubricroceus XxX
Notropis scabriceps
Notropis scepticus
Notropis semperasper
Notropis shumardi X XxX
Notropis spectrunculus

spectrunculus xX
Notropis spectrunculus subsp. xX X=
Notropis spilopterus

spilopterus
Notropis stilbius

x x
xx KK XK
x x

No Nx N § NeeN

x «x x
*

x KKK KKK KK
~

*
*

IP N N

x x

Notropis stramineus stramineus
Notropis telescopus
Notropis umbratilus

cyanocephalus XxX
Notropis volucellus
Notropis whipplei
Notropis sp. (paleband shiner)
Notropis sp. (sawfin shiner)
Notropis sp. (longirostris

group) xX
Phenacobius crassilabrum XxX
Phenacobius mirabilis Xx
Phenacobius teretulus
Phenacobius uranops
Phoxinus cumberlandensis
Phoxinus eos
Phoxinus erythrogaster
Phoxinus o. oreas
Phoxinus o. subsp.
Pimephales notatus
Pimephales promelas x
Pimephales vigilax XxX
Rhinichthys atratulus
Rhinichthys cataractae
Semotilus atromaculatus
Semotilus corporalis
Semotilus lumbee N N
Semotilus margarita
Tinca tinca

xx x x
*
~
~*~

N N N NI IP

x KKK XK
~
x «KKK XK

x KK
*
x KKK XK

IP N N NN eo eNeeIe Sse

»
OK XE Xi
|
J
HZ
HZ

x x >
x
rg

ING IN NG INE IN

> > > dO OOM
Med i
TAL VA
PATA TAA
ZZ
PANERA
ez ee

*
*
~
= Z

CATOSTOMIDAE

Carpiodes carpio Xx XxX
Carpiodes cyprinus XOX Xx N N N N N

Ss

VOLUME 95, NUMBER 1 35

Table 1.—Continued.

Drainage occurrence

Ohio Basin
Z i = z Native
2 & ae
2 i: ee
[) a pe) fas} o mo) ao) (oe)
aoe; ae Sp. eS > ise dae Mee eeu ae
see ee ee ee ae ee 2 gr Vere
ee es es Se ier aes ee ee ee age eee
OR eae eS ver vase ety SOT Bovey sa eh AG a 2
> ete ute RSs ICO EET EA RS) MOU Oy < za
Notropis photogenis ING NC ONT) UNE ND ON’ ONL EN” ZN ON: N O
Notropis procne procne No
Notropis p. longiceps IP So
Notropis pyrrhomelas So
Notropis rubellus Ne NS Ne N N N N*N N Ne Ne eNe eNoseeM
Notropis rubricroceus IP N So
Notropis scabriceps E
Notropis scepticus So
Notropis semperasper
Notropis shumardi N NP GM
Notropis spectrunculus
spectrunculus E
Notropis spectrunculus subsp. E
Notropis spilopterus
spilopterus NEN ONE NON N: N ON GNU UN WN, 2Nieg, NOM
Notropis stilbius N GM
Notropis stramineus stramineus Ni UN. UNG ON: IN’ oN. UN: ON N N M
Notropis telescopus Ieee N M
Notropis umbratilus
cyanocephalus N NN, ON oN ON- ON N M
Notropis volucellus NOON =Ne UN UN ON ON: ON) FN EN. GN eN GM
Notropis whipplei N N IN; ANG INP AN UN?) AN Nie N GM
Notropis sp. (paleband shiner) N N
Notropis sp. (sawfin shiner) N N
Notropis sp. (longirostris
group) N M
Phenacobius crassilabrum E
Phenacobius mirabilis N NN NTN 9 N] N N GM
Phenacobius teretulus E
Phenacobius uranops N N N
Phoxinus cumberlandensis Ja. 7-18
Phoxinus eos No M
Phoxinus erythrogaster N Ny aN UN NT UN] eNe ON UN M
Phoxinus o. oreas N IP
Phoxinus o. subsp. E
Pimephales notatus Ne NS UNT EN ON ON? OND ON ENN UN NP Pe Nowwem™
Pimephales promelas I i TPs. ff NN NON ON | NING S| NortGM
Pimephales vigilax N N NOON EN UN? UN NPN GM
Rhinichthys atratulus N NS ON N NN N NN NO NN NoSoF mM
Rhinichthys cataractae N N N N NoSo O
Semotilus atromaculatus NEN Ne ON NOON ON NON N NN (NOENoSoUGM

Semotilus corporalis No
Semotilus lumbee

Semotilus margarita N No M
Tinca tinca

CATOSTOMIDAE
Carpiodes carpio NS Ne Ne SIN] ON N GM
Carpiodes cyprinus NNN NEN NG EN] NON N NoSo GM

36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Drainage occurrence

Habitat Atlantic Slope

Lowland
Upland
Montane

Big River
Stream
Creek
Peedee

Cape Fear
Neuse

Tar

Roanoke
James

York
Rappahannock
Potomac
Susquehanna

x
~
Z

-Carpiodes velifer
Catostomus catostomus
Catostomus commersoni
Cycleptus elongatus
Erimyzon o. oblongus
Erimyzon o. claviformis
Erimyzon sucetta
Hypentelium etowanum
Hypentelium nigricans
Hypentelium roanokense
Ictiobus bubalus Xx
Ictiobus cyprinellus x
Ictiobus niger XxX
Lagochila lacera
Minytrema melanops xX
Moxostoma anisurum XxX
Moxostoma ariommum
Moxostoma atripinne
Moxostoma carinatum xX
Moxostoma cervinum
Moxostoma duquesnei
Moxostoma erythrurum
Moxostoma hamiltoni
Moxostoma m. macrolepidotum
Moxostoma m. breviceps
Moxostoma pappillosum
Moxostoma rhothoecum
Moxostoma robustum
Moxostoma rupiscartes

ICTALURIDAE

Ictalurus brunneus
Ictalurus catus
Ictalurus furcatus
Ictalurus melas
Ictalurus natalis
Ictalurus nebulosus
Ictalurus platycephalus
Ictalurus punctatus
Noturus baileyi
Noturus elegans
Noturus eleutherus
Noturus exilis

Noturus flavipinnis
Noturus flavus
Noturus furiosus x
Noturus gilberti
Noturus gyrinus
Noturus insignis

Noturus leptacanthus
es

x xX
x xX
x x
Zz,
Z
Z
Z,
Z,

Ne iN’ Ne aNeN

XxX «Ne N*IN: N- NPN ON @SNieNeN

x KK XK
xxx KKK XK
*
xx KKK

x
xX N N N N: N@eNeeNe ene aN
Xx E

IP

x KX

ZZ
Lae
iz,
ZA
mM Z

xx XK

x KK MK KK KK KK KK OK OK
~
x x xX ~

*
KKK mK KK KK KKK KK KK

~<
Za

x «x «x «x X
><
x «x x
><
Spd vA Vd edz
Z
Z
seed Ard
Z Z
Z Z
HZ
Z
Zag Z

x x XK
BP2Z2Z2Zz
Z

IP JP IP IPP sneer

xm KKK KK KK KK mK OK
*

x > x
>
Dede Dd DK DK DK Dd Dd Dd DK DK DK Dd De Dt Dx
Zaz
ZZ
Zaz
Zaz,
Pa 74 Va
ZzZZ
ZZ,

VOLUME 95, NUMBER 1

Table 1.—Continued.

37

Drainage occurrence

Ohio Basin
3 ie 5 E Native
a & 22 CGecutien
awe Ee Bo :
coe ee eo 5
ae Se CN sce Se eR > Soy 88e 08 A ee
cee Se ee ee ae oe Ss 5 ¢ 2 &
See ee Se Se Se Se ee aoe ee ER ops 0 Se ae
ONS Suite aver ce oe OO TO oy Pe Se ai oS = &
Zp ie ite Mie Oe ae Se IME Se (On Om ie < za
Carpiodes velifer N N ING Ne OND OUNE. UNE N, N So GM
Catostomus catostomus N No GM
Catostomus commersoni N N N N N N N N N N N N NoSo M
Cycleptus elongatus NP NP NP NP N NP N N N GM
Erimyzon o. oblongus NoSo
Erimyzon o. claviformis N N N GM
Erimyzon sucetta N So GM
Hypentelium etowanum NI G
Hypentelium nigricans NN NN N NON N NO’ N NN ENoSoNGM
Hypentelium roanokense
Ictiobus bubalus N N NON NEN UN WN N GM
Ictiobus cyprinellus N N N N N N GM
Ictiobus niger N NESNES NS IN N GM
Lagochila lacera N NP N N M
. Minytrema melanops N N NC -N) SND UN UN, ON NON So GM
Moxostoma anisurum N NP N N. No N UN - Ne ON NAN So M
Moxostoma ariommum
Moxostoma atripinne E
Moxostoma carinatum N N N NN -N INO Ne N N GM
Moxostoma cervinum
Moxostoma duquesnei N N N NG UNE SN UN, EN aN ae Ni GM
Moxostoma erythrurum IND eN SNe SSE TNT ENG ON Ne NING Ne eN GM
Moxostoma hamiltoni
Moxostoma m. macrolepidotum NoSo M
Moxostoma m. breviceps NE NSN Ne NE ONS SN] UNG SN N O
Moxostoma pappillosum So
Moxostoma rhothoecum N
Moxostoma robustum So
Moxostoma rupiscartes So G
ICTALURIDAE
Ictalurus brunneus So G
Ictalurus catus NoSo G
Ictalurus furcatus N N N N N N N GM
Ictalurus melas N N IP ND INS ENS ND END ONG EN GM
Ictalurus natalis N N N NI N N N N N N N N NOoSo GM
Ictalurus nebulosus N N NI N N N N IP N NoSo GM
Ictalurus platycephalus So
Ictalurus punctatus Ne NSN NIN Ne NSN] IN NS NEN So GM
Noturus baileyi E
Noturus elegans N N O
Noturus eleutherus No ONe UN OND ON N M
Noturus exilis N N N N M
Noturus flavipinnis E
Noturus flavus N N NEN NN NG Nn eN N M
Noturus furiosus
Noturus gilberti
Noturus gyrinus NNN N NoSo GM
Noturus insignis IP N IP NoSo
Noturus leptacanthus NI So G

38

Table 1.—Continued.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Lowland

Habitat

Montane
Big River

Stream

Creek

Drainage occurrence

Atlantic Slope

Peedee
Cape Fear
Neuse

Tar
Roanoke

James

York
Rappahannock
Potomac
Susquehanna

Noturus miurus
Noturus nocturnus
Noturus stigmosus
Noturus stanauli

Noturus sp. cf. leptacanthus

Pylodictis olivaris

AMBLYOPSIDAE

Amblyopsis spelaea
Chologaster agassizi
Chologaster cornuta
Typhlichthys subterraneus

APHREDODERIDAE

Aphredoderus sayanus

PERCOPSIDAE

Percopsis omiscomaycus

GADIDAE

Lota lota

CY PRINODONTIDAE

Fundulus albolineatus
Fundulus catenatus
Fundulus d. diaphanus
Fundulus heteroclitus
Fundulus lineolatus
Fundulus notatus
Fundulus olivaceus
Fundulus rathbuni
Fundulus stellifer
Fundulus waccamensis
Fundulus sp.

POECILIIDAE

Gambusia a. affinis
Gambusia a. holbrooki
Heterandria formosa

ATHERINIDAE

Labidesthes sicculus
Menidia extensa

GASTEROSTEIDAE

Apeltes quadracus
Culaea inconstans

COTTIDAE

Cottus baileyi
Cottus b. bairdi

~ xX

x KK XM

x KK

x «x XK

x KK KK

x x

x KKK

xX

XxX
XxX
Xx

Ma Ma Ma Ma Ma
Ma Ma Ma Ma Ma
N N N N N

Ma Ma Ma Ma Ma
NP N

NI

Ma Ma Ma Ma Ma
Ma Ma Ma Ma Ma

Ma Ma Ma Ma

Ma Ma Ma IP
IP

VOLUME 95, NUMBER 1 39

Table 1.—Continued.

Drainage occurrence

Ohio Basin
| = Native
Ss & & extralimital
& & Ss distribution
3 2 Oo a)
s s oS ae as a
Oo S a) 3 oO no} "e aS)
Be ee eS as > gs a 8
a ee Sa a cee ae ge aie le cae ees
See Bae SS OS See ben Jer eg: SeecleS
o 6k a 3 an ee) oO ¢ 5 5 5 = 2
ie Sy OMe EM tS) a ES aN SS Oy) a < a]
Noturus miurus N N N N -Ne oN UN ON. ON? NTN GM
Noturus nocturnus N N N N N GM
Noturus stigmosus N Ni SNE NF =e N: M
Noturus stanauli E
Noturus sp. cf. leptacanthus
Pylodictis olivaris Ne, ONY ON® ON: ONS ON: ON GN: UN] MINETIENE WAN GM
AMBLYOPSIDAE
Amblyopsis spelaea N O
Chologaster agassizi N N N M
Chologaster cornuta So
Typhlichthys subterraneus N N N GM
APHREDODERIDAE
Aphredoderus sayanus N N N NoSo GM
PERCOPSIDAE
Percopsis omiscomaycus Nei N UN N N N N No M
GADIDAE
Lota lota NI NI No M
CY PRINODONTIDAE
Fundulus albolineatus E
Fundulus catenatus N N N M
Fundulus d. diaphanus I No
Fundulus heteroclitus
Fundulus lineolatus So G
Fundulus notatus IN| IN| INT INT N GM
Fundulus olivaceus N N GM
Fundulus rathbuni So
Fundulus stellifer IP G
Fundulus waccamensis
Fundulus sp. N N
POECILIIDAE
Gambusia a. affinis IP IP Ma Ma IP Ma GM
Gambusia a. holbrooki NoSo G
Heterandria formosa So G
ATHERINIDAE
Labidesthes sicculus N N N I N N N N N N N N So GM
Menidia extensa
GASTEROSTEIDAE
Apeltes quadracus
Culaea inconstans I
COTTIDAE
Cottus baileyi E

Cottus b. bairdi NN NaN ENE NS UNG EN N N No M

40 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Drainage occurrence

Atlantic Slope

en)
ist)
a
=
ped}
oo

Lowland
Upland
Montane

Big River
Stream
Creek
Peedee

Cape Fear
Neuse
Roanoke
James

York
Rappahannock
Potomac
Susquehanna

Tar

Cottus bairdi subsp.
Cottus c. carolinae
Cottus carolinae subsp.
Cottus cognatus
Cottus girardi

Cottus sp. (smoky sculpin) Xx
Cottus sp. E

PERCICHTHYIDAE

Morone americana
Morone chrysops
Morone mississippiensis
Morone saxatilis

CENTRARCHIDAE

Acantharcus pomotis
Ambloplites cavifrons Xx
Ambloplites r. rupestris en XN EX
Centrarchus macropterus
Elassoma evergladei
Elassoma zonatum
Elassoma sp.

Elassoma sp.
Enneacanthus chaetodon
Enneacanthus gloriosus
Enneacanthus obesus
Lepomis auritus

Lepomis cyanellus
Lepomis gibbosus
Lepomis gulosus

Lepomis humilis

Lepomis macrochirus
Lepomis marginatus
Lepomis megalotis
Lepomis microlophus
Lepomis punctatus
Micropterus coosae
Micropterus d. dolomieui
Micropterus p. punctulatus
Micropterus s. salmoides
Pomoxis annularis
Pomoxis nigromaculatus

PERCIDAE

Ammocrypta asprella

Ammocrypta clara

Ammocrypta pellucida

Ammocrypta vivax xX
Etheostoma acuticeps

Etheostoma asprigene XxX

xx «KK
x KK KK
x KK KK

ZZ,

Ma Ma Ma Ma Ma Ma Ma Ma Ma Ma
I I I

xx x x
x KK XK
x KX

Ma Ma Ma Ma Ma Ma Ma Ma Ma Ma

*
*
Pd
Z
Za,
Z

NN Ne UN

IP br area aT

x KK KK
Z2ZZ5
ZZZ

Z Z
Zi Z,
Le Lie Za eZ,

x KKK KX KX
>
ZA ZZe AZ
Vd WA Na VD Wa Vey 7
ZZ ZZ ZZ
DID IZ 2 ZZ,
Z Diva 2 Zane
DD i ZZ
ZL ad,
ZZ4ZEZz
BAe ZZZ
Z,

x xX xX
xx x KK
ZZ
ZZ
Z5
ZH
5
5
cs
cs
5
ty Z22ZHzZ

x x

NG NEI iP I ot

x xX
x
mK K mK KKK KKK KK

x xX xX
xxx KX
xx x x
xxx KX

Ze

Zz,

Z,

Z

—
Ll
—_

Zaz
—_
—
—_
—

IP 1 ie eee

xx KK XK
x KKK XK
x KKK mK

VOLUME 95, NUMBER 1

Table 1.—Continued.

Drainage occurrence

4]

Ohio Basin
ez) 2) = = Native
ao as FS re extralimital
> ‘is iS) is distribution
ES ES ie 3a Te
Sig, oo eat eae 2 ¢ é
oes ee Wet Gee ee > SS elm Comey (2
Sy uN ME ee a ge Gen iS i es ae £2) on ee
oe 80 a a a wa os = 5 2 24 2 e ES
Se cae ee en Cae Came Ome oie cag caoey eee
Pree Ge a OD ASS PE MRS BO TO) Ge < za
Cottus bairdi subsp. N N
Cottus c. carolinae NON SN N M
Cottus carolinae subsp. N N
Cottus cognatus No
Cottus girardi
Cottus sp. (smoky sculpin) N So
Cottus sp.
PERCICHTHYIDAE
Morone americana NoSo
Morone chrysops I NI NI IP INT N Pai Nee Nie) N GM
Morone mississippiensis N N GM
Morone saxatilis I I I I I I NoSo G
CENTRARCHIDAE
- Acantharcus pomotis NoSo G
Ambloplites cavifrons
Ambloplites r. rupestris IND IN SOND CERT IN IN} ING? ONG ON] SNR NS NT} BINom eM
Centrarchus macropterus N N N So M
Elassoma evergladei So
Elassoma zonatum N N So GM
Elassoma sp. E
Enneacanthus chaetodon NoSo
Enneacanthus gloriosus NoSo G
Enneacanthus obesus NoSo
Lepomis auritus IP I I I I NoSo G
Lepomis cyanellus Ni OND SUNG) IPSN) ONE NE Ne) INT ING NON GM
Lepomis gibbosus NI NIP IP IP IP NoSo M
Lepomis gulosus NIP NN INS NG Se IN Som Givi
Lepomis humilis te N N N N NON GM
Lepomis macrochirus INS ING? NPS SINS ING IN) ON) © ONG NaN EN, So GM
Lepomis marginatus N So GM
Lepomis megalotis ING NING MT uN INS BING INI TINGS ING ee IN GM
Lepomis microlophus IP [Pee NEN N So GM
Lepomis punctatus N N So GM
Micropterus coosae I G
Micropterus d. dolomieui Ne ONG UNG TP? NY ON i DINO ONO ONG) FING goNfPr SiN! M
Micropterus p. punctulatus INN, ENG INTIS IN| OND Nite Ny ONS Ne eNP IN M
Micropterus s. salmoides Ne Ne ONY aR” UN: UN ON” UN) ON)’ UN) ee NIPSN So GM
Pomoxis annularis NY INN IP NU SINY ING, UN IN: GN ANTS N GM
Pomoxis nigromaculatus NON EN DP N N N N N N So GM
PERCIDAE
Ammocrypta asprella NN GM
Ammocrypta clara ING N N GM
Ammocrypta pellucida N’ ONY UN Ne Ne aNaeN) SN EN M
Ammocrypta vivax N GM
Etheostoma acuticeps E
Etheostoma asprigene NigEN N GM

42 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Drainage occurrence

Atlantic Slope

Habitat

Lowland
Upland
Montane

Big River
Stream
Creek
Peedee

Cape Fear
Neuse
Roanoke
James

York
Rappahannock
Potomac
Susquehanna

Tar

Etheostoma aquali
Etheostoma atripinne
Etheostoma barbouri
Etheostoma bellum
Etheostoma blennioides
blennioides
Etheostoma b. gutselli
Etheostoma b. newmanii
Etheostoma b.: newmanii X
blennioides
Etheostoma b.: n. X gutselli
Etheostoma blennius
Etheostoma boschungi
Etheostoma caeruleum xX
Etheostoma camurum xX
Etheostoma chlorobranchium xX xX
X
Xx

x eM KKK XK
xx x
xx KK
x KX

~
x
*

NI

x KK XK

Etheostoma chlorosomum xX
Etheostoma cinereum
Etheostoma collis collis XxX
Etheostoma c. lepidinion xX
Etheostoma duryi
Etheostoma etnieri
Etheostoma flabellare
Etheostoma fusiforme fusiforme
Etheostoma f. barratti
Etheostoma gracile
Etheostoma histrio
Etheostoma jessiae
Etheostoma kanawhae
Etheostoma kennicotti
Etheostoma longimanum
Etheostoma luteovinctum
Etheostoma m. maculatum
Etheostoma m. sanguifluum
Etheostoma m. vulneratum
Etheostoma mariae xX
Etheostoma meadiae
Etheostoma microlepidum
Etheostoma neopterum x
Etheostoma n. nigrum
Etheostoma n. susanae
Etheostoma n.: nigrum X

susanae
Etheostoma obeyense
Etheostoma olivaceum
Etheostoma o. olmstedi xX
Etheostoma o.: o. X

atromaculatum N
Etheostoma o. atromaculatum xX X Xe 2X IN| INE INP INP INT IN

x KK
xx KKK
x x xX
xm mK KK KKK XK
Za
Z,
Z,
Z,

x x

KKK KKK Km KK KK

N “Ne END aN

Km KKK KKK mK KK KK OK

x
x x x

xx KX
x x

N NN “N@N

VOLUME 95, NUMBER 1

Table 1.—Continued.

43

Drainage occurrence

Ohio Basin

Kanawha: below falls
Kanawha: above falls

Monongahela
Little Kanawha
Guyandotte

Big Sandy
Licking

Kentucky

Green

Cumberland: below falls

Cumberland: above falls

Tennessee

Native
extralimital
distribution

Atlantic Slope
Elsewhere

Etheostoma aquali
Etheostoma atripinne
Etheostoma barbouri
Etheostoma bellum
Etheostoma blennioides
blennioides
Etheostoma b. gutselli
Etheostoma b. newmanii
Etheostoma b.: newmanii x
blennioides
Etheostoma b.: n. X gutselli
Etheostoma blennius
Etheostoma boschungi
Etheostoma caeruleum
‘Etheostoma camurum
Etheostoma chlorobranchium
Etheostoma chlorosomum
Etheostoma cinereum
Etheostoma collis collis
Etheostoma c. lepidinion
Etheostoma duryi
Etheostoma etnieri
Etheostoma flabellare
Etheostoma fusiforme fusiforme
Etheostoma f. barratti
Etheostoma gracile
Etheostoma histrio
Etheostoma jessiae
Etheostoma kanawhae
Etheostoma kennicotti
Etheostoma longimanum
Etheostoma luteovinctum
Etheostoma m. maculatum
Etheostoma m. sanguifluum
Etheostoma m. vulneratum
Etheostoma mariae
Etheostoma meadiae
Etheostoma microlepidum
Etheostoma neopterum
Etheostoma n. nigrum
Etheostoma n. susanae
Etheostoma n.: nigrum X
susanae
Etheostoma obeyense
Etheostoma olivaceum
Etheostoma o. olmstedi
Etheostoma o.: 0. X
atromaculatum
Etheostoma o. atromaculatum

N°: AN GN ON, ON, No ON

ZZ,
ZZ,
ZZ,

NO UND ENG oN” oN, Ne ON

Ne NENG NG UN oN UN

esies

ZZ,

ZZ

Zm Zw

74, 74 \o| 74 74 Jeal teal leo

Os

GM

So

No GM

oT PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Drainage occurrence

Atlantic Slope

Habitat

Lowland
Upland
Montane

Big River
Stream
Creek
Peedee

Cape Fear
Neuse

Tar

Roanoke
James

York
Rappahannock
Potomac
Susquehanna

Etheostoma o.: o. X vexillare
Etheostoma o. vexillare
Etheostoma o. maculaticeps Xx
Etheostoma osburni
Etheostoma parvipinne xX
Etheostoma perlongum xX E
Etheostoma podostemone
Etheostoma proeliare Xx
Etheostoma rufilineatum
Etheostoma s. sagitta
Etheostoma sagitta spilotum
Etheostoma sellare
Etheostoma serriferum x
Etheostoma simoterum
Etheostoma smithi
Etheostoma s. spectabile
Etheostoma squamiceps
Etheostoma striatulum
Etheostoma stigmaeum
Etheostoma swaini XxX
Etheostoma swannanoa xX
Etheostoma tippecanoe
Etheostoma tuscumbia
Etheostoma variatum
Etheostoma virgatum
Etheostoma vitreum
Etheostoma z. zonale
Etheostoma sp. (duskytail darter) XxX X
Etheostoma sp. (Elk darter)
Etheostoma (Ulocentra) sp.
A—emerald darter
Etheostoma (Ulocentra) sp. B
Etheostoma (Ulocentra) sp. C
Etheostoma (Ulocentra) sp.
D—golden snubnose darter X xX OK
Etheostoma (Ulocentra) sp.
E—(Green River)
Etheostoma (Ulocentra) sp.
F—(Barren River) splendid
darter
Perca flavescens XxX
Percina aurantiaca
Percina burtoni
Percina c. caprodes
Percina c. semifasciata
Percina copelandi
Percina crassa
Percina e. evides
Percina evides subsp. XxX

Za
Zz

N- N

x KKK KK
x KK x

x KK
xx KX

N NN Nie N

xx KK KK
x KKK KKK KX
~< KK KK

x mK mK

x Kx x

N -N N N GNNeN

x KK KK
~
x Km

x
*

x
*
x x

x
~
*

NI NI NI NI NI NI NI NI NI NI

x KKK KK XX
~
x x xX

KKK KK KK XK
Z
Z

VOLUME 95, NUMBER 1 45

Table 1.—Continued.

Drainage occurrence
Ohio Basin
Native

extralimital
distribution

Kanawha: below falls
Kanawha: above falls
Cumberland: below falls
Cumberland: above falls

Monongahela
Little Kanawha
Guyandotte
Big Sandy
Licking
Kentucky
Tennessee
Atlantic Slope
Elsewhere

Green

Etheostoma o.: 0. X vexillare
Etheostoma o. vexillare
Etheostoma o. maculaticeps So
Etheostoma osburni E
Etheostoma parvipinne N
Etheostoma perlongum
Etheostoma podostemone
Etheostoma proeliare N
Etheostoma rufilineatum N N
Etheostoma s. sagitta E
Etheostoma sagitta spilotum E
Etheostoma sellare
Etheostoma serriferum So
Etheostoma simoterum NI NI
Etheostoma smithi
Etheostoma s. spectabile NY OEN ON:
Etheostoma squamiceps N
Etheostoma striatulum
Etheostoma stigmaeum N N
Etheostoma swaini
Etheostoma swannanoa
Etheostoma tippecanoe N N NN N' 'N
Etheostoma tuscumbia
Etheostoma variatum Ne ON N N N N N O
Etheostoma virgatum 18,
Etheostoma vitreum No
Etheostoma z. zonale NGS INGeN INGE INGOUN) NG ENE ON
Etheostoma sp. (duskytail darter) N
Etheostoma sp. (Elk darter)
Etheostoma (Ulocentra) sp. N N N
A—emerald darter
Etheostoma (Ulocentra) sp. B N
Etheostoma (Ulocentra) sp. C
Etheostoma (Ulocentra) sp.
D—golden snubnose darter N N
Etheostoma (Ulocentra) sp.
E—(Green River) E
Etheostoma (Ulocentra) sp.
F—(Barren River) splendid E
darter
Perca flavescens NI ree We
Percina aurantiaca
Percina burtoni N
Percina c. caprodes Nae NMaINS ANE NG ON NEDIN SEN) <iN
Percina c. semifasciata No
Percina copelandi N NON IN IND INGEN N
Percina crassa So
Percina e. evides N NN ee NF) NL aN N
Percina evides subsp. E
I ne ee ee he a

ZZz
ors

GM
GM

Ze t ZeZ to ZeZ ZZ,

eal 74 4

Fz,
= o

No

472 foal
<

SB Ss

46 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Drainage occurrence

Atlantic Slope

Habitat ~ =

E 2 5 . E a4 o 2 oe 5 ea = 2 3
Be a: BS es ith Bie Ta RB es oh a
a pS) Rae Ot ab OZ ee 2S eee

Percina gymnocephala x. EX X

Percina macrocephala Xx xX xX

Percina maculata x > ia. <

Percina n. notogramma x Xx Ne Ne aN

Percina n. montuosa x x E

Percina ouachitae x Xx Xo xX

Percina oxyrhyncha xX UX PS

Percina p. peltata Xx 4 NN OSNY SIN SN

Percina p. nevisense Xx XxX Ni“ .N; =N

Percina peltata subsp. Xx x E

Percina phoxocephala Xx Xx

Percina rex Xx Xx E

Percina roanoka x x NG N= IN Ni

Percina s. sciera Xr eX MK IN

Percina shumardi DS DS XxX

Percina squamata 2h DK Xx

Percina tanasi x xX

Percina (Odontopholis) sp. XxX Xx

Stizostedion canadense Xr OX x

Stizostedion v. vitreum XxX Xee GX! IP NI NI NI Ih IN

SCIAENIDAE
Aplodinotus grunniens Xe XxX XxX

Literature Cited

Hambrick, P. S., C. H. Hocutt, M. T. Masnik, and J. H. Wilson. 1973. Additions to the West Virginia
ichthyofauna with comments on the distribution of other species.—Proceedings of the West
Virginia Academy of Sciences 45:58—60.

Hendricks, M. L., J. R. Stauffer, Jr., C. H. Hocutt, and C. R. Gilbert. 1979. A preliminary checklist
of the fishes of the Youghiogheny River.—Natural History Miscellanea 203:1-15.

Hocutt, C. H., R. F. Denoncourt, and J. R. Stauffer, Jr. 1978. Fishes of the Greenbrier River, West
Virginia, with drainage history of the southern Appalachians.—Journal of Biogeography
5:59-80.

——_., , and . 1979. Fishes of the Gauley River, West Virginia.—Brimleyana 1:47-80.

, and P. S. Hambrick. 1973. Hybridization between the darters Percina crassa roanoka and

Percina oxyrhyncha (Percidae, Etheostomatini), with comments on the distribution of Percina

crassa roanoka in New River.—American Midland Naturalist 90:397—405.

; , and M. T. Masnik. 1973. Rotenone methods in a large river system.—Archiv fur

Hydrobiologia 72:245—252.

Jenkins, R. E., E. A. Lachner, and F. J. Schwartz. 1972. Fishes of the central Appalachian drainages:
their distribution and dispersal, pp. 43-117. In: P. C. Holt (ed.). The distributional history of
the biota of the southern appalachians. Part III: Vertebrates.—Research Division Monographs
4, Virginia Polytechnic Institute and State University. Blacksburg, Virginia.

VOLUME 95, NUMBER 1 47

Table 1.—Continued.

Drainage occurrence

Ohio Basin
= = Native
= S & = extralimital
Se 2 > distribution
Et Sela o «68
3 = 3 5 a < 2.
oR a) es} o as) as) 2
= peat ake Mey > pa Agari oO a 2
oD vy =| a Eo] = an Ts 5 5 A — r
5 o 2 2 = w S 2 5 2 2 2 = es
Be i= S > if ® = = I 5 es
Oe ee 3 a | wo 88 ) 2 5 =| O s 2
Siete ee Os PE Se a OE eee. ie < ea]
Percina gymnocephala E
Percina macrocephala N N N NPN UN N N O
Percina maculata N N N NPN NG NNN ON oN GM
Percina n. notogramma No
Percina n. montuosa
Percina ouachitae N N GM
Percina oxyrhyncha INN ONE SUN ENS NY ONS SIN? N
Percina p. peltata No
Percina p. nevisense
Percina peltata subsp.
Percina phoxocephala Nee Ne) N M
Percina rex
Percina roanoka IP
Percina s. sciera N N Ne Ne NE Ne NN N GM
Percina shumardi ING Ne NaeN N GM
Percina squamata N N
Percina tanasi E
Percina (Odontopholis) sp. N N
Stizostedion canadense N N N N NP N N UN N GM
Stizostedion v. vitreum N INN NiN| Nie NIE NN N GM

SCIAENIDAE
Aplodinotus grunniens N N N ING SuNi DAN INE SNL UN: N GM

Lee, D. S., C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J. R. Stauffer, Jr. 1980.
Atlas of North American freshwater fishes.—North Carolina State Museum of Natural History,
Raleigh, N. C. 854 pp.

Stauffer, J. R., Jr., K. L. Dickson, J. Cairns, Jr., and D. C. Cherry. 1976. The potential and realized
influences of temperature on the distribution of fishes in the New River, Glen Lyn, Virginia.—
Wildlife Monographs 50: 1-40.

—, C. H. Hocutt, and D. S. Lee. 1978. The zoogeography of the freshwater fishes of the Potomac
River basin, pp. 44-54. In: K. C. Flynn and W. T. Mason (eds.). The freshwater Potomac:
aquatic communities and environmental stresses.—Interstate Commission Potomac River Ba-
sin. Rockville, Maryland.

——,, , M. T. Masnik, and J. E. Reed, Jr. 1975. The longitudinal distribution of the fishes
of the East River, West Virginia-Virginia.—Virginia Journal of Science 26:121-125.

(JRS) Appalachian Environmental Laboratory, Center for Environmental and
Estuarine Studies, University of Maryland, Frostburg State College Campus,
Frostburg, Maryland 21532; (BMB) Department of Zoology, Southern Illinois
University, Carbondale, Illinois 62901; (REJ) Department of Biology, Roanoke
College, Salem, Virginia 24153; (CHH) Horn Point Environmental Laboratories,
Center for Environmental and Estuarine Studies, University of Maryland, Cam-
bridge, Maryland 21613.

PROC. BIOL. SOC. WASH.
95(1), 1982, pp. 48-57

TWO NEW GENERA OF DEEP-SEA POLYCHAETE WORMS
OF THE FAMILY AMPHARETIDAE AND THE ROLE OF
ONE SPECIES IN DEEP-SEA ECOSYSTEMS

Robert Zottoli

Abstract.—Two new ampharetid genera, Decemunciger and Endecamera, each
with one new species, D. apalea and E. palea, are described from wood panels
placed on the deep-sea floor by Turner (1973). The role of Decemunciger in deep-
sea ecosystems is discussed.

Recently, I examined a collection of ampharetid polychaetes removed from
pieces of wood collected by R. D. Turner, using the submersible DSRV Alvin,
from four experimental bottom stations in the North Atlantic, at depths ranging
from 1830 to 3995 meters. The wood had been placed at these sites by Turner to
study molluscan wood borers, and to “‘test the hypothesis that wood is an im-
portant source of nutrients and contributes to diversity in the deep sea’’ (Turner
1977:18).

Bivalve molluscs of the subfamily Xylophagainae (Family Pholadidae, Genera
Xyloredo and Xylophaga) mechanically excavate burrows in wood (Turner 1973,
1977). The bivalves ingest wood particles, making wood by-products available
within fecal pellets as food for detritus consumers. Adult and juvenile bivalves
may be consumed by a variety of predators such as galatheid crabs. Turner (1977)
suggests that young galatheid crabs may feed on recently settled Xylophaga lar-
vae and, later, on larger invertebrates including adult borers. Stomach contents
of older crabs contained sponge spicules, small pieces of wood, a nematode and
polychaete setae (Turner 1977). Other groups in this deep-sea food chain include
several families of polychaete worms, brittlestars, small sea urchins and predatory
gastropods (Turner 1977).

Two new genera from the family Ampharetidae, each with one new species,
have been found among the ampharetids associated with Turner’s wood panels.
The external anatomy of these ampharetids, Decemunciger apalea n. sp. and
Endecamera palea n. sp., is described, followed by a discussion of the role of
the former species in deep-sea ecosystems.

Materials and Methods

Three experimental islands, each with 12 separate one foot cubes of spruce
wood were placed by Turner (1977), for a period of five years, at the following
locations:

1. Deep Ocean Station 1 (DOS-1), 39°46’N, 70°41’W, 110 miles south of Woods
Hole, Mass., in 1830 m.

2. Deep Ocean Station 2 (DOS-2), 38°18.4’N, 69°35.6’W, 190 miles southeast of
Woods Hole, Mass., in 3506 m.

VOLUME 95, NUMBER 1 49

3. Tongue of the Ocean, Bahama Islands (TOTO Tower 3), 24°53.2'N,
77°40.2'W, in 2066 m.

Each experimental island is encircled by wood panels, 24” x 5” x 1”, which
are removed and replaced each time the islands are visited. At the time of re-
trieval, panels are enclosed in mesh bags to prevent loss of crumbled wood and
specimens. The mesh bags and their contents are then placed in retrieval boxes,
carried on the DSRV Alvin basket. The contents of the bags may be preserved
at the time the boxes are closed for return to the surface by puncturing plastic
bags previously placed in the retrieval boxes, thus releasing gluteraldehyde. Al-
ternatively, the contents in certain cases may be preserved immediately upon
reaching the surface. Specimens were well preserved. The use of submersibles
in studies of benthic communities is described by Grassle (1980a).

Systematics

Ampharetid polychaetes generally are wide anteriorly, tapering gradually to-
wards the posterior end (Fig. 1A). The prostomium is generally trilobed. Seg-
ments | and 2, which lie immediately behind the prostomium, are fused in most
species, and ventrally form the lower lip. Segment 3, in some species, bears one
lateral bundle of paleal setae on each side (Fig. 2A). The thorax begins at segment
4. The above segmental numbering system is that of Malmgren (1865-1866) and
Fauvel (1927), who recognized two segments in front of the paleal.

Decemunciger, new genus

Type-species.—Decemunciger apalea n. sp. Gender, masculine.

Diagnosis.—Body short, of 13 thoracic setigerous segments, last 10 unciniger-
ous, and of 14 abdominal uncinigerous segments. Segments | and 2 fused ventrally
forming lower lip. Segment 3 lacking paleae. Prostomium lacking glandular ridges.
Smooth, ventrally grooved, oral tentacles. Four pairs of smooth branchiae on
dorsal surface of segments 3-5. Branchial groups separated mid-dorsally by nar-
row space. Abdominal notopodia and notopodial and neuropodial cirri absent.

Remarks.—In comparison with other ampharetid genera with a similar distri-
bution of uncinigerous thoracic segments, Decemunciger differs from Melinnata
(Hartman, 1965), Melinnopsides (Day, 1964) and Muggoides (Hartman, 1965),
among other characters, by having 4 rather than 3 pairs of branchiae; from Mel-
innopsis (McIntosh, 1885) by lacking a fleshy ridge across dorsal surface of seg-
ment 6; and from Mexamage (Fauchald, 1972) by having 4 pairs of branchiae
inserted on 3 successive segments rather than 4.

Etymology.—Generic name derived from the Latin, refers to number of tho-
racic uncinigerous segments.

Decemunciger apalea, new species
Fig. 1A—C
Material examined.—(asb = asbestos-backed panel; D = DSRV Alvin).

Description.—Maximum size 6.3 mm long, 0.9 mm wide. Sexually mature in-
dividuals as small as 3.6 mm long, 0.54 mm wide. Holotype complete, 4.7 mm

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table of Material examined:

No. of specimens

Time
Panel Submerged Removed submerged Adult Juveniles
DOS-1
N28 8-30-75(D597) 9-28-77(D794) 2 yrs ) 1
N34 (asb) 8-30-75(D597) 7-29-77(D773) 2 yrs 2 32
N35 8-30-75(D597) 9-18-78(D834) 3 yrs 2 4
N47 6-15-76(D658) 7-30-77(D774) l yr 0 3
N65 8-17-76(D685) 7-29-77(D773) | yr 0 13
N67 8-17-76(D685) 8- 1-77(D776) 1 yr 0 7
N72 7-29-77(D773) 9-18-78(D834) 1 yr 0 3
N91 (asb) 9-28-77(D794) 9-18-78(D834) | yr ] 1
DOS-2
N31 9- 5-75(D601) 8- 3-77(D777) 2 yrs 16 10
Holotype (USNM #71545) and 4 paratypes (USNM #71546)
TOTO TOWER 3
T56 (asb) 5-12-77(D755) 11-11-78(D851) 1 yr 0 1

long, 0.9 mm wide. Color in alcohol white to pale orange. Prostomium indistinctly
trilobed, lacking glandular ridges. About 14 smooth oral tentacles, each with deep
ventral groove. Segments | and 2 fused, ventral part forming lower lip. Segment
3 without paleae and not visually obvious. Four pairs of smooth branchiae, about
’% body length; 2 on segment 3, 1 on segment 4, and | on segment 5. Branchial
groups separated mid-dorsally by narrow space. Notopodial lobes each bearing
7-11 winged capillary setae from segments 4—16. Each seta about 0.4 mm long,
7.5 wm wide basally, and 10 wm wide across the blade. Notopodia lobes (unci-
nigerous pinnules) bearing toothed uncini from segment 7 to end of abdomen.
Ten thoracic and 14 uncinigerous abdominal segments. Thoracic uncini in single
transverse rows, 22-35 per row. Each with about 10 teeth, more or less in 3
transverse rows, above a rounded basal prow (Fig. 1B). Abdominal uncini in
single tranverse rows, about 10-18 per row. Each with about 15 teeth in several
transverse rows, above a rounded basal prow (Fig. 1C). Abdominal notopodia
and notopodial, neuropodial, and anal cirri lacking. Pygidium rounded.

Remarks.—Mucus-lined tubes covered with particulate matter, about 3 times
worm length. Female about 4 mm long and 0.5 mm wide with approximately 260
elliptical eggs in body cavity ranging from 25 to 150 wm across widest diameter.
No gonoducts visible.

Etymology.—Specific name, derived from the Latin, refers to lack of paleae.

Endecamera, new genus

Type-species.—Endecamera palea n. sp. Gender, feminine.

Diagnosis.—Body short, of 14 thoracic setigerous segments, last 11 unciniger-
ous and of 14 abdominal uncinigerous segments. Segments | and 2 fused, ventrally
forming lower lip. Paleae present on segment 3. Prostomium lacking glandular
ridges. Smooth, ventrally grooved, oral tentacles. Four pairs of smooth branchiae

VOLUME 95, NUMBER 1 51

A

Fig. 1. Decemunciger apalea: A, Lateral view of entire worm, 5.2 mm long; B, Mid-thoracic
uncini, lateral and frontal views, length = 16 wm; C, Mid-abdominal uncini, lateral and frontal views,
length = 14 um.

a2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

on dorsal surface of segments 3-4. No mid-dorsal space between branchial groups.
Abdominal notopodia and notopodial and neuropodial cirri absent.

Remarks.—In comparison with other ampharetid genera with a similar distri-
bution of thoracic uncinigerous segments, Endecamera differs from Ampharana
Hartman, 1967; Amythasides Eliason, 1955, and Glyphanostomum Levinsen,
1884, by having 4 rather than 3 pairs of branchiae; from Amage Maimgren,
1865-1866, Grubianella McIntosh, 1885, and Phyllampharete Hartman and Fau-
chald, 1971, by lack of abdominal notopodia; from Pterampharete Augener, 1918,
and Sabellides Milne Edwards in Malmgren, 1865-1866, by having smooth rather
than papillose oral tentacles, and from Paramage Caullery, 1944, by notosetae
beginning on segment 4 rather than segment 6.

Etymology.—Generic name, derived from the Greek, and transcribed to the
Latin with a feminine-singular ending, refers to number of thoracic uncinigerous
segments.

Endecamera palea, new species
Fig. 2A—C

Material examined.—St. Croix Station, 17°57.63'’N, 64°48.6’W, in 3995 m
DSRV Alvin dive 876, 20 Dec 1978, ‘‘wild’’ wood about 6 feet long. Eighteen
specimens. Holotype (USNM 71547); 3 paratypes (USNM 71548).

Description.—Maximum size 5 mm long, 0.75 mm wide. Holotype complete,
3.3 mm X 0.5 mm. Color in alcohol white to pale orange. Trilobed prostomium
with middle lobe about same width as lateral lobes; lacking glandular ridges.
Smooth, ventrally grooved, oral tentacles. Segments 1 and 2 fused ventrally form-
ing lower lip. Two lateral groups of paleae, about 11 in each group on segment
3. Each palea approximately 0.38 mm long, 5 wm wide basally, tapering gradually
to a fine point. Four pairs smooth branchiae, about %4 body length; 3 of each
group in a straight line across dorsal surface of segments 3-4 with 4th inserted
just anterior to most medial branchiae. No mid-dorsal gap between branchial
groups. Notopodial lobes, each bearing 7-12 winged capillary setae from segment
4 to end of thorax. Setae about 0.43 mm long, 8 wm wide basally, and 10 um
wide across the blade. Notopodia of segment 4 minute while those of segments
5 and 6 larger than those of segment 4, but smaller than those of subsequent
segments. Fourteen thoracic setigerous segments. Neuropodial lobes bearing
toothed uncini from segment 7 to end of abdomen; 11 thoracic and 14 uncinigerous
abdominal segments. Thoracic uncini in single transverse rows, about 23-27 per
row. Each with 10 teeth, more or less in 2 transverse rows, above a rounded
basal prow (Fig. 2B). Abdominal uncini in single transverse rows, about 10-18
per row. Each with about 13 teeth in several rows above a rounded basal prow
(Fig. 2C). Abdominal notopodia and notopodial, neuropodial and anal cirri lack-
ing. Pygidium rounded.

Remarks.—Mucus-lined tubes covered with particulate matter, about 3 times
body length. Sexually mature females were broken or twisted making it impos-
sible to count the number of eggs in the body cavity. No gonoducts visible.

Etymology.—Specific name, derived from the Latin, refers to presence of pa-
leae on segment 3.

VOLUME 95, NUMBER 1 53

Fig. 2. Endecamera palea: A, Lateral view of entire worm, 4 mm long; B, Mid-thoracic uncini,
lateral and frontal views, length = 14 wm; C, Mid-abdominal uncini, lateral and frontal views,
length = 12 um.

54 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Feeding of the Two Species

Ampharetid polychaetes live in mucus-lined tubes covered by particulate mat-
ter. The tube often extends some distance above the sediment surface. Jumars
found that starved Hobsonia increased the length of their tubes more, over a
period of several months, than those that were fed (Fauchald and Jumars 1979).
They suggested that tube building may be a form of locomotion, allowing animals
to enter new feeding areas. Fauchald and Jumars (1979) also suggested that hor-
izontal or vertical tube orientation may depend on food availability; horizontal
tube orientation would allow worms to cover a larger feeding area.

During feeding, worms emerge from the tube opening and evert their oral ten-
tacles equal to about their body length over the sediment surface (Fauchauld and
Jumars 1979; personal observations on Asabellides oculata [Webster], Hobsonia
[Amphicteis| floridus [Hartmen] and Melinna cristata [Sars]). Food adheres to
mucus produced on the ventral side of the tentacles and is carried by cilia to the
mouth. The tentacular apparatus of Decemunciger apalea and Endecamera palea
appears similar to that of shallow water ampharetids and it is assumed that the
method of feeding is also similar. Pieces of wood, detritus and bivalve larvae
(Xylophaga sp.) were found in the gut cavity of Decemunciger apalea, while only
detritus was noted in the digestive tract of Endecamera palea.

The digestive tract of both species like most ampharetids consists of buccal
cavity, oesophagus, stomach, intestine, and rectum. The buccal cavity houses
about 14 ventrally grooved and ciliated, smooth oral tentacles.

Food available to deep-sea organisms includes plankton, remnants of marine
macrophytes such as Sargassum, remnants of land plants, particulate residues
of deteriorating nekton and chemo-autotrophic bacteria found in hydrothermal
vent areas (George and Higgins 1979, Rowe and Staresinic 1979). Rowe and
Staresinic (1979), using sediment traps, found that about 4 g C/m?/yr, mainly in
the form of fecal pellets, reached the deep-sea bottom. Marine macrophyte re-
mains were next in importance. Pieces of the brown alga Sargassum accounted
for an average of 0.4 g C/m?/yr while the contribution from other sources was
relatively small. |

Decemunciger apalea most likely feeds on fecal pellets produced by other
animals on the “wood islands,”’ on bivalve larvae (Xylophaga sp.) that settle
near them, and on any type of detrital particle that reaches them from the sur-
rounding water column. Fungi and bacteria on small pieces of ingested wood may
represent an added source of nutrients.

Life History/Recruitment/Growth Rate of Decemuniger

The rate of colonization in deep-sea sediments is about two orders of magnitude
lower than in shallow water (Grassle 1977). In addition, larval recruitment and
settlement, growth rates, and probably mortality rates, are generally lower in the
deep sea (Grassle and Sanders 1973, Grassle 1977, Sanders 1979). Grassle (1980b)
found fewer individuals and species in boxes of azoic sediment placed on the
deep-sea floor, compared to samples taken from surrounding sediments. Because
of these factors and others, deep-sea populations are commonly dominated by
mature adults (Grassle 1977). Opportunistic wood boring, deep-sea bivalves
(Subfamily Xylophagainae; Family Pholadidae) characterized by rapid growth,

VOLUME 95, NUMBER 1 55

Table 1.—Numbers of Decemunciger apalea (in parentheses) of various body lengths (in mm) for
listed wood panels.

N31 DOS 2: (1) 2.0; (4) 2.7; (3) 3.0; (2) 3.2; (5) 3.6; (1) 4.0; (7) 4.5; (1) 5.4; C1) 6.0; (1) 6.3.

N32 DOS 1: (2) 0.4; (1) 0.6; (1) 0.65; (1) 0.85; (5) 1.0; (2) 1.2; (3) 1.4; (2) 1.75; C1) 1.8; C1) 1.9; (2)
e038 (C)) 232 (8) e523 (1) 252 (Ob) S02 Gly S552 (10) S05

N34 and

NOS DOS me (0! 752 (1) 028; (2): 1:0; (2) 1.25; GB) 1.53 C1) 1:75; (1) 1.75: CG) 2.5; GC) 3:0.

ING DOSE) th2--() 153: (2) 1.5; CU) 1.75; (1) 2.5; C1) 2:65.

ING2Z DOS 1: (1) 1.5; (1) 3.0; (1) 4.0; (1) 4.5; C1) 35.25.

N91 DOS 1: (1) 1.0; (1) 4.5.

early sexual maturity, and relatively large numbers of eggs, appear to be an
exception to the generality above (Turner 1973, 1977). The larger number of eggs
and the resultant large number of motile larvae allow at least a few individuals
to reach geographically scattered pieces of wood (Turner 1973, Sanders 1979).

Decemunciger appears to have a life cycle similar to these bivalves. The large
number of small eggs in the body cavity implies a high reproductive rate. The
apparent onset of sexual maturity within a year’s time, documented below, sug-
gests early maturity and a rapid growth rate. Thus, also considering its ability to
colonize a transient habitat, Decemunciger may be characterized as an oppor-
tunistic species. Evidence for this is as follows, the only unknown aspect being
the method of larval dispersal.

Decemunciger with well developed egg or sperm in the body cavity ranged
from about 3.6 to 6.3 mm in total body length. Worms less than 3.6 mm long were
therefore assumed to be juveniles. Eggs probably pass singly through certain
nephridia and nephridiopores into the anterior part of the tube where they are
fertilized by sperm, released by males in the same fashion. Hobsonia (Amphic-
teis) floridus (Zottoli, 1974), Hypaniola kowalewski (Marinescu, 1964) and Mel-
innexis artica (Annekova, 1931), which have eggs roughly the same diameter as
Decemunciger, retain developing larvae in the maternal tube until they are able
to crawl on the bottom. It is hypothesized that these brooding and colonization
patterns are the same for Decemuniciger.

Sexually mature worms as well as juveniles were found on wood panels N72
and N91 which were submerged for one year. Thus, if one accepts the premise
that wood panels are colonized solely by larvae, then sexual maturity is attained
within a year (Table 1). The absence of sexually mature Decemunciger on wood
panels N34, N65, and N67, which were also submerged for one year, suggests
that larval worms settled on the wood late in the year and did not have sufficient
time to reach sexual maturity. In addition, the presence of certain size classes of
adults and juveniles on wood panels N31, N34, N72, and N91 suggests that this
species reproduces only at certain times of the year. Rokop (1974) felt that deep-
sea organisms generally reproduce throughout the year. Lightfoot, Tyler, and
Gage (1979) suggest that cyclic seasonal breeding is more common in the deep-
sea than previously supposed. Timing of reproduction most likely reflects sea-
sonal abundance of food reaching the deep-sea floor (Lightfoot, Tyler, and Gage
1979).

56 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Acknowledgments

Thanks are due to Dr. J. Fred Grassle, Charlene D. Long, Dr. Meredith L.
Jones, and Dr. Ruth D. Turner for making specimens available and for reviewing
the manuscript. The study, conducted by Turner (1977), is supported by the
Office of Naval Research (ONR Contract No. 14-76-C-1281, NR 104-687 to Har-
vard University). This paper is dedicated to my recently departed friend and
colleague, Dr. Fred (Eric) Davis.

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Sanders, H. L. 1979. Evolutionary ecology and life-history patterns in the deep sea.—Sarsia 64: 1-7.

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Fitchburg State College, Fitchburg, Massachusetts 01420.

PROC. BIOL. SOC. WASH.
95(1), 1982, pp. 58-66

THE KARYOTYPE OF THE EURASIAN FLYING SQUIRREL,
PTEROMYS VOLANS (L.), WITH A CONSIDERATION OF
KARYOTYPIC AND OTHER DISTINCTIONS IN
GLAUCOMYS SPP. (RODENTIA: SCIURIDAE)

V.R. Rausch and R. L. Rausch

Abstract.—The karyotype of Pteromys volans (L.) Qn = 38; FN = 73) is de-
scribed, based on material from Hokkaido and northeastern Siberia, and com-
pared with those of Glaucomys volans (L.) (2n = 48; FN = 80) and Glaucomys
sabrinus (Shaw) (2n = 48; FN = 78) from North America. Although the similar-
ities of the fundamental numbers are suggestive of karyotypic evolution of the
Robertsonian type, the fossil record of these flying squirrels is at present frag-
mentary, and no conclusion is drawn as to their affinities. Definite differences
were observed between the karyotypes of G. volans and G. sabrinus, which had
been described as identical. In combination with karyotypic differences, other
characteristics of the two North American species indicate both a divergence
earlier and an age greater than have been considered on the basis of paleontologic
evidence. Karyograms of the three species of flying squirrels are included.

In the course of investigations concerning northern mammals, we studied chro-
mosomal preparations from two specimens of the Eurasian flying squirrel, Ptero-
mys volans (L.), from Hokkaido and northeastern Siberia. Chromosomes of the
two species of flying squirrels of the Nearctic genus Glaucomys were compared.
The purpose of the present report is to define the karyotype of P. volans and to
describe some previously unrecognized differences in the karyotypes of the two
Nearctic species. Other distinguishing characteristics of Glaucomys spp. also are
briefly discussed.

Materials and Methods

Two male specimens of P. volans were utilized. The first, P. volans orii Ku-
roda, was provided in 1967 by Dr. H. Abe, who captured the animal at Koshimizu,
Abashiri, Hokkaido Island (ca. 43°58’N, 144°30’E). Cells from bone marrow and
from one testis were fixed and stained in acetic orcein, following standard meth-
ods for mammalian karyology. The second, P. volans cf. incanus Miller, was
trapped by us in August 1979 along the upper Kolyma River, Magadansk Oblast’,
near the settlement of Sibik-Tiellakh (ca. 62°N, 149°30’E). Cells from marrow
(femur) were prepared in the field and processed with Giemsa blood stain (Sea-
bright 1972). In both cases, cells were treated with colchicine.

Preparations were made by the same methods for Nearctic flying squirrels, as
follows: Glaucomys sabrinus yukonensis (Osgood), two males collected at An-
chorage and near Fairbanks, Alaska, in October 1966 and April 1969, respective-
ly; one male G. sabrinus bangsi (Rhoads), captured on Powwatka Ridge, Wallowa
Mountains, Oregon (ca. 45°40’N, 117°27’W), in December 1980; and one male G.
volans querceti (Bangs), collected in the vicinity of Tampa, Florida, in early 1980.

VOLUME 95, NUMBER 1 59

After chromosomes (in metaphase) were counted, representative cells were
photographed on high-contrast film, and the component chromosomes in five cells
from each animal were measured (excepting the squirrels from the upper Kolyma
River and from Anchorage, in which only contracted chromosomes were ob-
served in the preparations). Chromosomes exhibiting Giemsa-banding were seg-
regated according to size and banding-pattern with use of photographs and light
microscopy. The identification of the X-chromosome was based on measurements
and pattern of banding. The Y-chromosome in Glaucomys spp. was selected
subjectively, since the small submetacentric chromosomes in these animals, of
which the Y is one, appeared to be similar in both size and banding. The termi-
nology concerning the location of the centromere on individual chromosomes
follows Levan et al. (1964). The number of major chromosomal arms (funda-
mental number, or FN) was determined according to the procedure of Matthey
(1945). The chromosomal components of 30 to 61 cells from each of the animals
were counted, excepting those from the upper Kolyma River and Anchorage for
which only 18 and 15 cells, respectively, were counted.

Results

I. Pteromys volans (2n = 38, n = 19).—The diploid complement was made up
of 12 metacentric autosomes (arm-ratios ranged from 1.03 to 1.7), 22 submeta-
centric-subtelocentric and 2 acrocentric autosomes (arm-ratios from 2.4 to 6.3),
the submetacentric X-chromosome (arm-ratio from 1.6 to 1.8), and the acrocentric
Y-chromosome (arm-ratio from 4.2 to 7.3). In 61 cells (53 mitotic and 8 meiotic)
examined, 49 contained 38 chromosomes and 7 contained 19 bivalent elements.
A karyogram from the male P. volans from Hokkaido is shown in Fig. 1.

The set of chromosomes (in all of the 18 cells examined) from the animal
collected in northeastern Siberia was morphologically like that of P. volans from
Hokkaido, consisting of 6 pairs of metacentric and 12 pairs of submetacentric-
acrocentric autosomes, a submetacentric X, and an acrocentric Y-chromosome,
as shown in Fig. 2.

The number of major autosomal arms was 70 in both animals. Two autosomes
with thin and lightly stained short arms were considered to be acrocentric. The
FN (autosomes + X and Y) was 73 for P. volans.

II. Glaucomys spp. (2n = 46).—Karyograms from G. volans (Florida) and from
G. sabrinus (Oregon and Alaska) are shown in Figs. 3—5. Thirty bi-armed (meta-
centric-subtelocentric) chromosomes (with arm-ratios ranging from 1.03 to 8.8)
and 16 acrocentric chromosomes (with centromeres in the terminal region; arm-
ratios ranged from 4.5 to 13.8) constituted the autosomal set in G. volans. In G.
sabrinus, from both localities, the autosomes included 28 metacentric-subtelo-
centric (arm-ratios ranging from 1.07 to 8.6) and 18 acrocentric (arm-ratios from
4.3 to 14.5) elements. The sex-chromosomes of the two species were similar
morphologically and in pattern of banding: X submetacentric, of medium size
(ranges of arm-ratios were 1.5 to 2.03 for G. volans, and 1.9 to 2.9 for G. sa-
brinus), and Y submetacentric of small size (ranges of arm-ratios were 2.1 to 3.2
for G. volans, and 2.0 to 3.3 for G. sabrinus). The patterns of bands indicated
that these two species have some equivalent, or homologous, chromosomes.
However, significant morphologic differences also were evident.

60 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

6

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Figs. 1-5. Karyograms of Pteromys volans (2n = 38) and Glaucomys spp. (2n = 48). 1, P. volans,
male, from Hokkaido Island; Orcein stain. 2. P. volans, male, from the Magadansk Oblast; Giemsa
blood stain. 3, G. volans with Giemsa-bands, male, from Florida. Arrow indicates metacentric ele-
ments not present in G. sabrinus. 4, G. sabrinus with Giemsa-bands, male, from Oregon. Arrow
indicates acrocentric elements not present in G. volans. 5, G. sabrinus, male, from Alaska; Orcein
stain. Scale-lines have value of 5 wm.

VOLUME 95, NUMBER 1 61

Discussion

The gliding squirrels allocated to the genera Pteromys G. Cuvier, 1800, and
Glaucomys Thomas, 1908, have had rather complex taxonomic histories and, as
indicated by recent reviews (cf. Mein 1970, Black 1972), their origins and affinities
are obscure. The grounds for acceptance of the genus Pteromys instead of Sci-
uropterus F. Cuvier, 1825, for the flying squirrel in northern Eurasia have been
discussed by Miller (1914) and by Ellerman and Morrison-Scott (1951), among
others, but Simpson (1945) considered the latter to be probably the valid generic
designation. In addition to the widely occurring P. volans, the genus includes a
second species, P. momonga Temminck, 1846, which occurs on the Japanese
Islands of Honshu, Kyushu, and Hondo. According to Ellerman and Morrison-
Scott (1951), its cranial differences are so marked that it cannot be considered a
race of P. volans.

In his revision of the genera and subgenera of the Sciuropterus-group of flying
squirrels, Thomas (1908) recognized four subgenera in Sciuropterus, mainly on
the basis of dental characters. These included Sciuropterus, with S. (Sciuropter-
us) russicus Tiedemann, 1808 (=Sciurus volans Linnaeus, 1758) as type, and
Glaucomys, which was erected for the Nearctic flying squirrels, with S. (Glau-
comys) volans (=Mus volans Linnaeus, 1758) as type. Glaucomys was elevated
to generic rank by Howell (1915), on grounds not stated, but he later (1918)
defined cranial and dental characters that distinguish Glaucomys from Pteromys.
These distinctions were confirmed by Ognev (1940), who also pointed out the
marked differences in the structure of the os penis in P. volans and G. volans.
Recently, from the study of both fossil and living flying squirrels, Mein (1970)
placed Pteromys and Glaucomys in different generic groupings on the basis of
dental characters.

The fossil record provides no clear indication of the origin of Pteromys volans.
However, the comparison of serum proteins by means of double immunodiffusion
by Zholnerovskaia et al. (1980) suggested that Pteromys has its closest affinities
with Sciurus and Tamias, rather than with terrestrial sciurids. In North America,
Glaucomys is known from the middle to late Irvingtonian. The stratigraphic range
of Glaucomys volans extends from the late Irvingtonian, whereas remains of the
northern G. sabrinus are known only from deposits of late Wurm age (Kurtén
and Anderson 1980). Thenius (1972) considered that Glaucomys appears to have
arisen from terrestrial sciurids, which had their center of radiation in North
America.

The diploid number of 38 chromosomes, as reported by Tsuchiya (1979) and
determined by us, alone distinguishes the karyotype of Pteromys volans from
those of Glaucomys spp. (2n = 48) (Nadler and Sutton 1967, Schindler et al.
1973). In the former, 35 chromosomes were metacentric-subtelocentric, and only
3 (two autosomes and the Y-chromosome) were acrocentric. The relatively large,
acrocentric Y of P. volans is markedly different from the male sex-chromosomes
of Glaucomys spp. That the fundamental numbers in these squirrels are rather
similar (73 in P. volans; 78 in G. sabrinus; and 80 in G. volans) suggests that
karyotypic evolution involving chromosomal rearrangements of Robertsonian
type might have occurred. However, no significance is attributed to the apparent
similarities when nothing is known concerning the further relationships of the
Eurasian and American species.

62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

The karyotype of a female specimen of P. momonga has been published by
Tsuchiya (1979, fig. 7). It appears to be quite similar to that of P. volans, but the
available details do not permit an extensive comparison. The presence of boreal
forest far to the south in the Japanese Islands during the glacial maximum of
Wurm time would seem to have favored the southward dispersal of Pteromys
volans (cf. Kotani 1969).

With reference to the karyotypes of Glaucomys spp., the published information
is discrepant. That G. volans and G. sabrinus are karyotypically identical has
been generally accepted. However, for both, Schindler et al. (1973) considered
the number of bi-armed (metacentric-subtelocentric) autosomes to be two more
than did Nadler and Sutton (1967). Hsu and Benirschke (1973) concurred with
Schindler et al. The karyotype defined by us for G. sabrinus yukonensis (Alaska)
agreed more closely with that shown and discussed by Nadler and Sutton (1967),
who studied preparations from G. sabrinus lascivus (Bangs) (California), than
with the results of Schindler et al. (1973) for G. sabrinus from New Hampshire
(an area where this species is marginally sympatric with G. volans). Accordingly,
we examined new material from G. sabrinus as well as G. volans, from geographic
regions far removed from areas of parapatry or sympatry (Oregon and Florida).
The identities of the animals studied were confirmed from the distinctive char-
acteristics of the os penis, in addition to other criteria.

Although the respective karyotypes of these flying squirrels appeared superfi-
cially to be much alike, they exhibited distinct differences, principally in the
presence in G. volans of two small metacentric autosomes that did not occur in
G. sabrinus, and in G. sabrinus of two small acrocentric autosomes not observed
in G. volans. As well, the total lengths of individual chromosomes differed be-
tween the two species, including those in some presumed to be homologous; the
relative length of the female haploid complement (autosomes + X) was recorded
as 65 wm in G. volans and 86 um in G. sabrinus. Morphologically, the sex-
chromosomes were similar.

Of the chromosomes of G. volans, the two small metacentric autosomes (see
Fig. 3, arrow) were the most nearly metacentric, sensu stricto, with an arm-ratio
ranging from 1.03 to 1.05. However, these and the two small acrocentric elements
in G. sabrinus (see Fig. 4, arrow) were similar in total length; their value relative
to the lengths of the respective haploid complements was the same (3.5%) in both
species. The appearance of Giemsa-bands in these chromosomes suggests that
they may be homologues, and that the small metacentric pair in G. volans arose
as a result of a pericentric inversion involving the small acrocentric pair in G.
sabrinus (or perhaps vice versa). The disparate total lengths of the respective
complements further suggest quantitative differences in nuclear DNA, which
were not defined by our methods.

Our findings indicate that the two species of Glaucomys are karyotypically
distinct, and they are in contrast with the conclusions of others concerning these
flying squirrels. The karyogram from G. volans from Florida (Fig. 3) appears to
be identical with those shown by Schindler et al. (1973) as representative of both
G. volans and G. sabrinus in New Hampshire, where the two species are sym-
patric.

The two species of Glaucomys are distinguished additionally by major differ-
ences in the form of the os penis, as is also Pteromys volans. Since detailed

VOLUME 95, NUMBER 1 63

Table 1.—Helminths of Glaucomys volans and G. sabrinus.

G. sabrinus
G. volans

Midwest Midwest Oregon
(n = 22) (n = 5) (n = 26)
Species of helminth no. infected no. infected no. infected
Nematoda:
Capillaria americana Read, 1949 7 — —
Syphabulea thompsoni (Price, 1928) 1 5 a=
Syphabulea sp.* — — 17
Lemuricola sciuri (Cameron, 1932) 13 a a
Citellinema bifurcatum Hall, 1916 2 D y
Cestoda:
Andrya sciuri Rausch, 1947 — 2 4
Monoecocestus thomasi Rausch and Maser, 1977 — — 18
Catenotaenia sp. — 3 —
Hymenandrya sp. — — 3

* Description in preparation by J.-P. Hugot, Museum National d’Histoire Naturelle, Paris.

descriptions of these structures have been published, it suffices here to point out
their diversity in the three species. The os penis of P. volans is small (ca. 4 mm
long in our material) and consists of a slender, pointed bone with a well developed
barb at the proximal end (cf. Ognev 1940, fig. 153). That of G. volans, first
described by Pocock (1923), is long (ca. 14 mm in specimens from Florida) and
very slender, whereas that of G. sabrinus is shorter (ca. 6.5 mm in our material),
broader, and flattened (see Burt 1960, plate IV). Each differs markedly from the
others, and we were unable to discern any fundamental similarities unless, per-
haps, the barb near the distal end of the os penis of G. sabrinus is homologous
with the basal barb in that of P. volans.

Certain differences in the helminth faunas of the two species of Glaucomys
provide a further indication of dissimilarity. Of the nine species of helminths
recorded by us from flying squirrels in North America (Table 1), six occur also
in sciurids of other species (Rausch and Tiner 1948, Davidson 1976, McGee 1980).
Host-specific cestodes of three species, Andrya sciuri, Monoecocestus thomasi,
and Hymenandrya sp., are known only from Glaucomys sabrinus. The assem-
blages involving these cestodes would seem to have arisen independently through
coevolution of helminth and host (Rausch 1981), indicating comparatively early
divergence of the two species of Glaucomys. These cestodes are Nearctic species,
with no congeners known from Pteromys volans or other sciurids in Eurasia. The
only helminth known from P. volans is Citellina petrovi Shul’ts, 1930, which is
host-specific and has been recorded from European Russia, Japan, and Chukotka
(Hugot 1980). Some of the characteristic ectoparasites of Pteromys volans and
Glaucomys spp. are congeneric (Acarina and Mallophaga), but no species is
shared. Consequently, these arthropods do not appear to provide any useful
indications of the relationships of their hosts.

Various hypotheses have been proposed to account for the essential allopatry
of the Nearctic flying squirrels, but the geographic ranges of these mammals seem
simply to reflect differences in specific ecologic requirements. The two species
occupy disjunct ranges over the greater part of North America where they occur,

64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

relative to the distribution of coniferous and deciduous forest. In the east, where
their ranges are parapatric to sympatric, they appear as well to be segregated
ecologically. As noted by Guilday (1962), G. sabrinus has a relict distribution in
the Appalachian Mountains, where it occurs at elevations above ca. 1000 m as
a consequence of climatic change (warming) during post-glacial time.

That the characteristic habitats of the two species of Glaucomys have a fun-
damental relationship to their different dietary requirements is evident. Glauco-
mys volans utilizes buds, fruits, and insects during spring and summer, but later
feeds mainly on acorns and other nuts and seeds, some of which are stored in
large quantities and consumed during winter (cf. Muul 1968). Glaucomys sabrinus
depends mainly on hypogeous fungi during the warmer months, and utilizes epi-
phytic lichens (Usnea and Alectoria) during winter (McKeever 1960, Maser ef
al. 1978, Maser pers. comm.). The diet of Pteromys volans differs from these in
that it feeds on sprouts or buds of Betula and Acer and on male flowers of conifers
in summer, and on catkins or buds of Betula and Alnus in winter (Ognev 1940,
Tavrovskii et al. 1971).

Suggestions have been made to account for the pattern of distribution of Glau-
comys sabrinus on grounds other than ecologic requirements. Muul (1968) con-
sidered that the southward spread of G. sabrinus is prevented by competitive
interaction with G. volans, inasmuch as the latter breeds earlier seasonally and
therefore would already occupy the limited number of tree-cavities available.
However, as reported by Cowan (1936), and observed by us in northern Wiscon-
sin, G. sabrinus commonly utilizes outside nests constructed mainly of twigs.

Weigl (1975) observed in captive animals that infection by a nematode of the
genus Strongyloides was tolerated by G. volans, but was associated with high
mortality in G. sabrinus. He considered that differential pathogenicity of this
nematode might account in part for the segregation of the two species of Glau-
comys where they are sympatric. Barbehenn (1978) suggested that this might
constitute a mechanism by which the larger G. sabrinus would be excluded from
habitat otherwise suitable for G. volans. However, Strongyloides robustus Chan-
dler, 1942 exhibits little host-specificity, occurring in sciurids of various species
(Rausch and Tiner 1948, McGee 1980). Davidson (1976) considered it to be the
most pathogenic of the more common helminths in squirrels, and reported that
infections involving more than 150 individuals often caused severe enteritis in
gray squirrels, Sciurus carolinensis Gmelin. It might be expected that transmis-
sion of this nematode would be enhanced by conditions of captivity, and that
differences in feeding habits or other behavior could account for more massive
infections in G. sabrinus. That Weig] did not find northern flying squirrels natu-
rally infected by Strongyloides appears to be compatible with the results of other
surveys of helminths in this sciurid.

Altogether, the differences between the two flying squirrels of the genus
Glaucomys seem to indicate earlier divergence and greater age of these species
than has been considered on the basis of paleontologic evidence. The origins and
affinities of Glaucomys spp. and of Pteromys volans may be established if the
earlier fossil record of these sciurids can be traced.

Acknowledgments

Specimens of flying squirrels were kindly provided by H. Abe, Faculty of
Agriculture, Hokkaido University, Sapporo; Ms. E. L. Bull, Pacific Northwest

VOLUME 95, NUMBER 1 65

Range and Habitat Laboratory, U.S. Department of Agriculture, La Grande,
Oregon; R. Anderson, Wallowa Valley Ranger District, U.S. Department of Ag-
riculture, Joseph, Oregon; and I. L. Duncan, Center for Disease Control, Atlanta.
Field work in the Soviet Union was carried out under the U.S.A./U.S.S.R. En-
vironmental Protection Treaty, Area V, Subproject A-3. In the U.S.S.R., V. L.
Kontrimavichus, D. I. Berman, and Ms. A. N. Leirikh, Institute of Biological
Problems of the North, Academy of Sciences of the U.S.S.R., Magadan, kindly
provided support and assistance. We express sincere thanks for these contribu-
tions.

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Thenius, E. 1972. Grundziige der Verbreitungsgeschichte der Saugetiere. Eine historische Tiergeo-
graphie. Gustav Fischer Verlag, Jena. 345 pp.

Thomas, O. 1908. The genera and subgenera of the Sciuropterus group, with descriptions of three
new species.—The Annals and Magazine of Natural History (8)1:1-9.

Tsuchiya, K. 1979. A contribution to the chromosome study in Japanese mammals.—Proceedings of
the Japanese Academy (B)55:191-195.

Weigl, P. D. 1975. Parasitism as a possible biological weapon affecting the ranges and interactions
of the flying squirrels, Glaucomys volans and G. sabrinus. American Society of Mammalogists,
55th Annual Meeting, 16-19 June 1975. No. 36, Abstracts of Technical Papers, p. 17.

Zholnerovskaia, E. I., N. N. Vorontsov, and O. K. Baranov. 1980. Immunologicheskii analiz sis-
tematicheskikh vzaimootnoshenii letiag (Pteromys) c piat’iu rodami palearkticheskikh
belich’ikh (Sciuridae, Rodentia).—Zoologicheskii Zhurnal 59:750—754.

Burke Memorial Washington State Museum, DB-10, and Division of Animal
Medicine, SB-42, University of Washington, Seattle, Washington 98195.

PROC. BIOL. SOC. WASH.
95(1), 1982, pp. 67-80

A NEW SPECIES OF THE GENUS ECHININUS (MOLLUSCA:
LITTORINIDAE: ECHINININAE) WITH A
REVIEW OF THE SUBFAMILY

Joseph Rosewater

Abstract.—A new ovoviviparous species, Echininus viviparus, inhabiting high
intertidal and supratidal areas in the Mariana Islands is described in the littorinid
subfamily Echinininae which previously consisted of the polytypic Indo-Pacific
species Echininus cumingi (Philippi, 1846) and the west Atlantic species Tectin-
inus nodulosus (Pfeiffer, 1839). Characteristics of the subfamily are reviewed.

I received for identification from L. G. Eldredge of the University of Guam
specimens of an Echininus which upon close examination proved to be not only
a new species, but one which exhibits ovoviviparity. The species described here
I originally thought to be Echininus cumingi spinulosus (Philippi, 1847), and the
realization that it is not prompted me to take a fresh look at the subfamily Echin-
ininae resulting in the following review.

Family Littorinidae Gray, 1840
Subfamily Echinininae Rosewater, 1972

Diagnosis.—The subfamily Echinininae is characterized by spinose to bluntly
spinose shells; umbilicate or imperforate; opercula multispiral; radulae with mod-
erately to greatly narrowed rachidians. Distribution is tropical west Pacific and
west Atlantic.

The subfamily Echinininae was established by Rosewater (1972) to contain the
genus Echininus which differs in matters of sculpture, radula, animal morphology,
and ecology from either Littorininae or Tectariinae. This subfamily concept is
strengthened by findings enumerated in the present paper. Members of the
subfamily have a unique habitat among Littorinidae, for they inhabit highest shore
levels, at times seeming almost to be terrestrial. The radula of Echininus, while
said to be “‘not unusual’’ (Rosewater 1972:507), is distinctive when examined
with the SEM (Fig. 1A—D). The radula of Tectininus is unique (Fig. 1E—F). The
operculum is multispiral, a departure from all other known littorines. Character-
istics of both radulae and opercula are here considered adaptations to a high
shore habitat, as is the occurrence of ovoviviparity in the new species of Echin-
inus described herein. All are indicative of a basic evolutionary trend in Echini-
ninae toward a terrestrial habit. Shells of this group are also distinctive, especially
that of E. cumingi cumingi, which is characterized by rather long, open spines
and a deep umbilicus. These latter features, also, may be of significant adaptive
value to an animal living on tropical shores enabling easier dissipation of heat.

Tectininus Clench and Abbott, 1942

Tectininus Clench and Abbott, 1942:4; type-species by original designation Echin-
inus nodulosus (Pfeiffer, 1839).—Abbott, 1954:458-462.

68 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Diagnosis.—Shells bluntly spinose, succeeding spiral rows of spines non-syn-
chronous; imperforate; opercula multispiral; radula with very reduced rachidian,
modified lateral, and marginals with few denticles. Distribution is subtropical to
tropical west Atlantic.

As originally proposed by Clench and Abbott (1942) and elaborated by Abbott
(1954) the monotypic subgenus, Tectininus, was distinguished from Echininus
s.s. because its type, E. nodulosus, possesses a unique radula, although still
exhibiting the formula 2-1-1-1-2 and a modified littorinid embayment (Fig. 1E—F).
It is unlike any other radula in the family, having massive lateral teeth and a
diminutive rachidian which appears almost vestigial. The operculum is multispir-
al. It possesses a penis bearing a subterminal penial gland surrounded with fleshy
papillae, which differs from the condition in the nominate subgenus and combines
characteristics noted in the genera Littorina, Nodilittorina, and Tectarius. Aj-
though Rosewater (1972) had not seen the paper and reported that nothing was
known of reproduction in Echinininae, it had already been shown that 7. nodu-
losus 1S Oviparous and produces a pelagic egg capsule (Borkowski 1971).

Until further evidence for its true relationships comes to light, I am satisfied
to permit Tectininus to remain in its present position. As stated by Abbott, after
careful study of the type-species of Tectininus, it appears to be **. . . a special-
ization of the ancestral stock . . . inthe family Littorinidae’’ (Abbott 1954), whose
apparent closest relations are in the Echinininae. Because of its unique radula
Tectininus should be accorded full generic status.

Echininus Clench and Abbott, 1942

Echininus Clench and Abbott, 1942:3; type-species by original designation Tro-
chus cumingii Philippi, 1846.—Rosewater, 1972:525.

Diagnosis.—Shells spinose to bluntly spinose; often with partly open spines;
umbilicate to imperforate; opercula multispiral; radulae with moderately nar-
rowed rachidians, laterals and marginals with moderately numerous denticles.
Distribution is west Pacific from southern Japan through western Pacific arc to
Cook Islands. |

The genus Echininus was delineated as consisting of 2 subgenera containing a
total of 2 living species and | subspecies (Rosewater 1972). The nominate sub-
genus, Echininus, inhabits the west Pacific and contains E. cumingi cumingi
(Philippi, 1846) known from the southern Philippines southward through Mela-
nesia (including New Guinea, Solomon Islands, New Hebrides, New Caledonia)
eastward to the Cook Islands, and, E. cumingi spinulosus (Philippi, 1847) which
occurs in southern Japan, the Ryukyu Islands, Taiwan and northern Philippines.
Echininus adelaidensis (Cotton, 1947) is a Pliocene fossil from South Australia,
which resembles certain trochaceans (Rosewater 1972). New evidence from pre-
served specimens indicates that the Mariana Islands population, previously be-
lieved to be part of E. cumingi spinulosus, is different from either of the previ-
ously known Pacific subspecies and is a new species.

Echininus cumingi cumingi, type-species, is comparatively large, reaching 20.9
mm (Table 1), with 3 spiral rows of conspicuous projecting, open spines on the
body whorl; it has a narrow, deep umbilicus; a multispiral operculum is present;
the radula has a moderately narrow rachidian, lateral and inner marginal teeth

VOLUME 95, NUMBER 1 69

each having one large and several smaller cusps, and slender outer marginals
having 4—5 cusps. Echininus cumingi spinulosus is very close in appearance to
E. c. cumingi, although it never reaches as large a size (18.3 mm), is less spinose,
and has a narrower umbilicus which occasionally is closed. The specimens from
the Mariana Islands which were included with the former by Rosewater (1972)
are consistently different from either E. c. cumingi or E. c. spinulosus. Available
records indicate they are limited in distribution to the southern Mariana Islands,
and, so far have been reported only on the islands of Saipan, Tinian, Rota and
Guam. Specimens are smaller in size, the largest measuring 12.3 mm in length;
have only 2 major rows of blunt spines on the body whorl. In addition, the shell
shape is different; obesity of the Mariana specimens averages 0.76 compared with
0.87 in cumingi and 0.78 in spinulosus. There are a number of quite absolute
anatomical and biological differences also. Females of the Mariana population
reproduce ovoviviparously, and the expanded oviduct may contain large numbers
of young. The embryos are visible through the transparent dorsal mantle tissue
in various stages of development including young snails with shells of 1.25—1.5
whorls. This condition is very similar to that described in Littorina saxatilis by
Thorson (1946), and Fretter and Graham (1962). The reproductive modes of E.
c. cumingi and E. c. spinulosus are not known, but there is no evidence from
examination of preserved specimens to indicate Ovoviviparity in them. There are
also morphological differences between radulae and penes of the Mariana species
and the more western and northern Echininus.

Echininus viviparus, new species
Figs. 1-6, Table |

Description.—Shell: reaching 12.3 mm (about /% inch) in length, turbinate in
shape, with blunt spines; average obesity (width/length) about 0.76 (76 specimens
range from 0.65—0.84) relatively thick in structure, usually non-umbilicate or with
only a thin slit between columella and parietal callus; suture usually evident
although often partially obscured by spiral sculpture; whorls 3—6, moderately
shouldered below suture, relatively flat-sided; usually with 2 major rows of blunt
spines on body whorl and one on spire whorls; occasionally with one or more
additional major rows of spines. External shell color varying from grayish to
tannish orange, the darker color characteristic of shells retaining periostracum;
periostracum frequently worn away on spines which then appear grayish white.
Apertural coloration light to dark orange with occasional light or darker orange
lines revolving within. Base moderately flattened, sculptured with weakly nod-
ulose cords. Length of spire greater than half length of shell. Spire convex; spire
angle quite variable: from 60—72°. Aperture subquadrate; outer lip thickened,
smooth within, only slightly wavy at edge reflecting external sculpture; inner lip
with small but distinct toothlike bulge. Suture evident when not partly obscured
by sculpture. Primary sculptural feature: 2 spiral rows of irregular blunt spines
on body whorl, and one row on spire whorls. Rows of spines not well aligned
axially; body whorl spine count |1—16 in posterior row and 12—16 in anterior row;
13-14 on penultimate whorl of spire. Secondary spiral sculpture consists of 3-4
fat, non-spinose cords between each spinose row, which occasionally become
nearly as spinose as spinose cords they separate. In addition, an overall micro-

70 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

f ‘ 4 as : f ; ”
i a ; te Z.
| ‘ Ld i ath ‘ ie $ < 3

Fig. |. Radulae: A, B, Echininus viviparus, 2° from northeast coast of Tinian, Mariana Islands
(USNM 796244): A, Showing 2 complete transverse rows, Bar = 10 wm, 875; B, Rachidian tooth,
Bar = 5 um 1800. C, D, Echininus cumingi cumingi, 2 from Davao, Mindanao, Philippines (USNM
747765): C, Showing 2 complete transverse rows, Bar = 10 wm, 700; D, Rachidian tooth, Bar = 5
pm, 1400. E, F, Tectininus nodulosus, 2 from San Salvador, Bahamas (USNM 596683): E, Showing
1 complete transverse row; note small, narrow rachidian tooth and massive laterals, Bar = 10 wm,
510; F, Oblique view of rachidian tooth, Bar = 2 wm, 3500.

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VOLUME 95, NUMBER 1

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72 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Shells of Echininus viviparus: A, B, Holotype (USNM 792356), 9.7 x 7.4 mm; C, D,
Paratype (USNM 803296) 10.7 x 7.6 mm.

scopical sculpture of fine, closely-spaced threads. Axial sculpture consists of
irregular lines of growth. Operculum small in size, multispiral (polygyrous spiral
type of Fretter and Graham 1962) having S—6 volutions, chitinous, light brown in
color. Nuclear whorls usually present, smooth, light brown to reddish brown;
reaching about 1.25—1.50 volutions before developing spiral sculpture.

Animal: Radula littorinoid, 2-1-1-1-2; rachidian narrow, but with bulbous base
having 3 basal denticles; lateral tooth with large central cusp and smaller lateral
cusps and with distinct littorinoid embayment in which inner marginal tooth ar-

VOLUME 95, NUMBER | 73

WW OE

ww SZ

Fig. 3. Echininus viviparus: A, 2 with well developed young in expanded oviduct seen through
dorsal mantle; B, Detail from A: shell of young snail of about 1.5 whorls; C, d showing arrangement
of penial glands. All from northeast coast Tinian, Mariana Islands (USNM 796244).

ticulates; inner marginal with 4 cusps of which third from medial is largest; outer
marginal with 4—5 cusps increasing in size laterally. Animal littorinoid; in the
male penis is unbranched; wide and deep sperm groove runs along its posterior
surface, the edges of which are brown pigmented; penis with 5-8 penial glands
arranged in straight line along outer edge; well developed ctenidium present in

74 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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Fig. 4. Known distribution of Echininus species.

dorsal mantle cavity; leaflets well formed; efferent branchial vessel and osphra-
dium present (similar to that shown in Fretter and Graham 1962, Fig. 7). Fecal
pellets oval in shape, white with dark flecks. Female ovoviviparous with oviduct
modified as brood pouch; containing as many as 50 or more embryos in various
stages of development up to the crawling stage in which shell has 1.25-1.50
whorls.

Etymology.—Viviparus, an adjective, from the Latin, meaning ‘‘bearing active,
living young.”’ }

Measurements.—See Table 1.

Types.—Holotype, USNM 792356; length 9.7 mm, width 7.4 mm; |5 paratypes,
USNM 803296, ranging in length from 8.1—12.3 mm, and in width from 6.2—8.2
mm.

Type-locality.—Machong Point [=As Matmos or Dudu, personal communi-
cation, L. G. Eldredge] (about 14°11’N, 145°18’E), Rota, Mariana Islands; in
spray zone 30 feet landward from top of 50 foot cliff; 4 February 1979; L. G.
Eldredge, collector.

Material examined.—Mariana Islands, West Pacific Ocean.

Guam:
[1*] Agfayan Point (13°16'N, 144°44’E) high intertidal on limestone; July 1980,
R. H. Randall, USNM 803297.

* Figure ‘‘1’’ refers to column | in Measurements Table 1; specimens from Guam are summarized
together.

VOLUME 95, NUMBER 1 M5

Camel Rock (13°29’N, 144°42’E) at 20 foot elevation on limestone; 13
June 1980, L. G. Eldredge, USNM 803298.

Asan Point (13°29'N, 144°42’'E); 1951, D. B. Langford, USNM 613687.

Apra Bay (13°27'N, 144°38’E); November 1907, P. Bartsch, USNM
PBB323.,

Guam, 29 August 1949, V. L. Haack, USNM 620383.

Guam, 1946, Capt. Draeger, USNM 707190.

Rota:

[2] Machong Point [=As Matmos or Dudu] (about 14°11'N, 145°18’E) in spray
zone 30 feet landward from top of 50 foot cliff; 4 February 1979, L. G.
Eldredge, type-lot, USNM 792356, 803296.

[3] West side Poniya Point (14°06'N, 145°10’E) above terraced bench in spray
zone, 2 February 1979, R. K. Kropp, USNM 803299.

Tinian:

[4] Northeast coast (about 15°06’N, 145’39’E) in spray and splash zone of large
blowhole, 100 feet landward from 20 foot cliff, 8 February 1979, L. G.
Eldredge, USNM 796244.

Saipan:

[5] Puntan Agingan (15°07'N, 145°42’E) in depressions in limestone 6-10 feet
landward from 25-30 foot cliff, 19 November 1980, L. G. Eldredge,
USNM 803300.

[6] Puntan Magpi (15°16’N, 145°48’E) on limestone in spray of blowhole, 5-8
foot cliff, 21 November 1980, L. G. Eldredge, USNM 803301.

Ecology.—Lives in the high intertidal and supratidal areas on rough, pitted
limestone apparently restricted in distribution by the extent of ocean spray; often
in the vicinity of blowholes (Tinian) and sometimes found at considerable dis-
tances landward from the tops of sea cliffs 20-50 feet high (Rota and Tinian) (L.
G. Eldredge, personal communication) (Fig. 6).

Echininus viviparus lives highest above the sea of any littorinid that I have
observed. It is described as living where it is only wet by spray from surf or the
blowholes characteristic of coral shores, where it probably feeds on vegetation
which is kept moist by the same spray. The ovoviviparous reproduction of this
species appears to be in close accord with its high position on the shore and
permits it to survive without a pelagic stage in its development. Careful studies
need to be carried out before it can be stated unequivocally, however, that E.
viviparus has no need to return to the sea for any part of its life history.

Geographical distribution.—Southern Mariana Islands: Guam, Rota, Tinian,
and Saipan.

Comparative remarks.—Morphometrics of Echininus viviparus are compared
with those of E. c. cumingi and E. c. spinulosus in Table 1. Note that E. viviparus
is smaller than the other two species, reaching only 12.3 mm in maximum length
versus 20.9 and 18.3 mm respectively. On the average it is less obese, 0.76 versus
0.87 and 0.78, although difficulties of measuring spinose shells may artifically
enhance the obesity of E. c. cumingi. Numbers of postnuclear whorls and rows

76 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig.5. A, Apex of Echininus viviparus, showing smooth protoconch of about 1.25 whorls (compare
with Fig. 3B), specimen from northeast coast Tinian (USNM 796244), Bar = 100 um, 100x; B, Apex
of Littorina saxatilis, another ovoviviparous littorine, showing smooth protoconch of about 1.5
whorls, specimen from Odiorne Point State Park, Rye, Rockingham County, New Hampshire (USNM
803282), Bar = 100 wm, 80x; C, Apertural view of young E. viviparus showing 2 major rows of blunt
spines on body whorl separated by moderately strong intermediate row, same data as A, Bar = 250
pm, 40x; D, Apex of E. cumingi cumingi, showing decollate tip, specimen from Philippines (USNM
89449), Bar = 100 wm, 185x.

of tubercles are less in E. viviparus. The aperture length/shell length index is
smaller in E. viviparus. These data show the new species is smaller in size and
more slender than previously known species. It is also non-umbilicate where the
other two usually have an umbilicus. There is a rather weak but persistent tooth-
like bulge on the inner lip.

The animal of E. viviparus is typically littorinoid, having a blunt snout and
tentacles with eyes at the outer bases. It is significant that females of FE. viviparus
produce young ovoviviparously (Fig. 3A) because this has not been noted in other
Echininus species. The penis of E. viviparus differs markedly from other species:
in E. viviparus there are 5-8 penial glands in a straight line along the outer edge
of the unbranched penis (Fig. 3C), while in E. c. cumingi, the glands occur only

VOLUME 95, NUMBER 1 VI

at the junction of the bulbous base and the penial extremity, laterally and pos-
teriorly (Rosewater 1972: plate 406, fig. B). The arrangement in E. c. spinulosus is
similar to the latter. Radulae of the three species of the genus Echininus are
similar, with differences in some details. In all species the teeth are small (see
magnifications, Fig. 1) and reminiscent of the ‘‘pick type’’ (Rosewater 1980).
Rachidian teeth of all three species of Echininus are narrow distally, but in E.
viviparus the base is bulbous with 3 denticles (Fig. 1B). The tip of the protoconch
is frequently present in E. viviparus, but is smaller and often decollated in E.
cumingi and E. spinulosus (Fig. 5).

Discussion.—When | initially mapped the distribution of Echininus (Rosewater
1972, plate 407), information from museum specimens indicated that the ranges
of the two subspecies, FE. c. cumingi and E. c. spinulosus, were divided into
northern and southern components by an east-west line drawn just south of Puerto
Princessa, Palawan, Philippines (9°N). Some additional records have come to
light for E. c. cumingi: Guimaras Island, Philippines [verifying the type-locality
of E. cumingi] (10°35'N) (Kevin Marx, personal communication 1979); Lifou,
Loyalty Islands (C. Lamb 1979, USNM) not very far from the Tana, New Heb-
rides record reported by Rosewater (1972). The discovery of the new species,
Echininus viviparus, changes the pattern of distribution of the genus Echininus
in the Indo-Pacific (Fig. 4). Although obviously related generically, the Mariana
species is quite distinct from the other Echininus taxa in its distribution, in certain
aspects of its morphology and, so far as is known, in reproducing ovoviviparous-
ly. In my previous evaluation of a few small, worn shells from Guam (Rosewater
ibid.) I referred them to E. c. spinulosus because no anatomical evidence was
available to the contrary. The additional freshly collected specimens provided by
L. G. Eldredge have shown without doubt that E. viviparus is a separate species
distinct from the subspecies group of E. cumingi. It is rather difficult to speculate
upon the origin of FE. viviparus. Little fossil evidence exists to trace the evolu-
tionary history of the genus Echininus or, in fact, of any members of the family
Littorinidae. Some littorinid fossils may have been assigned to such families as
Turbinidae and Trochidae which are almost indistinguishable as fossils. The most
similar in appearance is the fossil, Tectarius rehderi Ladd, 1966, from Lower
Miocene, Marshall Islands (Ladd 1966, Rosewater 1972, pl. 404, figs. 5-7). Ac-
cording to Ladd (1960) much of the mollusk fauna now inhabiting west Pacific
islands and the East Indies may have migrated in successive waves from east to
west during Cretaceous and Tertiary times when an archipelago of islands existed
through which species could move with relative ease. Such species as Tectarius
rehderi may be the precursors of Recent Echininus and Tectarius species now
inhabiting the western Pacific arc. It seems unlikely that the fossil species Echin-
inus adelaidensis (Cotton, 1947) from the Pliocene of South Australia belongs in
the Littorinidae and probably should be referred to the Trochacea. In the absence
of data indicating a more widespread distribution, it appears that Echininus vi-
viparus has evolved as an endemic species in the Mariana Islands where its
ancestors became established during one of these earlier migrations. Other lit-
torinid inhabitants of the Mariana Islands fail to show any indication of such
endemicity (Roth 1976, Rosewater 1970, 1972).

Echininus viviparus joins the small but increasing group of mollusks whose life
histories are known to include ovoviviparity, or brooding of young by the parent

78 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ie
oe
Sa
is

Fig. 6. A, Machong Point, Rota Island, Mariana Islands, habitat of Echininus viviparus 30 feet
landward from top of 50 foot cliff; B, Same, showing spray reaching habitat; C, Same, showing pitted
limestone; white dots are Echininus; Bar = 20 cm; D, Northeast coast Tinian, Mariana Islands, where
Echininus lives 100 feet landward from top of 20 foot cliff. (Photos courtesy of L. G. Eldredge.)

through early stages of development. The occurrence of this form of reproduction
in mollusks was reviewed by Van der Schalie (1936) who attributed its develop-
ment to being of survival value for animals subjected to unfavorable environment.
Ovoviviparity now is known to occur in several additional groups to those cited
by Van der Schalie (1936). In Littorinidae it is known in at least three species,
L. saxatilis (=L. littoralis of Van der Schalie), and E. viviparus. Littorina scabra
scabra and L. scabra angulifera release young in various stages of development,
as summarized by Mileikovsky (1975). Other instances of ovoviviparity in gas-
tropods occur in Siliquariidae, Struthiolariidae, Nassariidae, and Turridae (Mor-
ton 1958); Coralliophilidae (Wells and Lalli 1972); Janthinidae and Hydrobiidae
(Fretter and Graham 1962); Acmaeidae, Hipponicidae, Planaxidae, Vermetidae,
Thiaridae, a few opisthobranchs, many Stylommatophora, a number of chitons,
and some Aplacorphora (Hyman 1967). Among the Bivalvia there are also scat-
tered cases: Nuculidae (Drew 1901); freshwater bivalves except Dreissensia [sic]
(Cooke 1895); Arcidae, Ostreidae, Erycinacea (Morton 1958); Carditacea (Jones
1963, Yonge 1969); Veneridae (personal observation in the genus Gemma); Ter-
edinidae (Turner 1966).

Ovoviviparity occurs within groups of mollusks often in what appears to be a
rather haphazard fashion, although occasionally it will occur in an entire family
such as Unionidae or Viviparidae. The development of this special type of pa-
rental care, as Van der Schalie (1936) has suggested, appears associated with
animals living under special stress for which ovoviviparity contributes survival

VOLUME 95, NUMBER 1 79

value, i.e. larvae of freshwater mussels are distributed on fish; crawl-away young
of Echininus living on cliffs far above the sea will survive where a pelagic de-
veloper could not. There is no way to predict when ovoviviparity will occur in
a group. But the frequency with which it does occur lends credence to its value
under the proper conditions.

Summary.—As presently defined the littorinid subfamily Echinininae consists
of the following units distributed as noted:

Family Littorinidae Gray, 1840
Subfamily Echinininae Rosewater, 1972
Genus Echininus Clench and Abbott, 1942, type-species by original desig-
nation Trochus cumingii Philippi, 1946
Echininus cumingi cumingi (Philippi, 1846)—southern Philippines and
along the western Pacific arc to New Hebrides and the Cook Islands
Echininus cumingi spinulosus (Philippi, 1847)—southern Japan through the
Ryukyu Islands to northern and western Philippines
Echinininus viviparus, new species—southern Mariana Islands
?¢Echininus adelaidensis (Cotton, 1947)—Pliocene, South Australia
Genus Tectininus Clench and Abbott, 1942, type-species by original desig-
nation, Litorina nodulosa Pfeiffer, 1839
Tectininus nodulosus (Pfeiffer, 1839)—southern Florida, Bermuda, the
Bahamas, and the Greater Antilles

Acknowledgments

L. G. Eldredge, University of Guam, provided specimens of Echininus vivip-
arus which he sent to me via G. J. Vermeij and P. Signor. He also sent habitat
photos and information on ecology. The female with brood pouch and details
of male anatomy were drawn by I. Jewett. SEM preparations were made by
P. Greenhall and scanning electron microscopy performed at the Smithsonian’s
SEM laboratory, under the direction of W. Brown, by S. Braden and M. Mann.
R. S. Houbrick offered helpful suggestions on the manuscript.

Literature Cited

Abbott, R. T. 1954. Review of the Atlantic periwinkles, Nodilittorina, Echininus and Tectarius.—
Proceedings of the U.S. National Museum 103(3328):449-464.

Borkowski, T. V. 1971. Reproduction and reproductive periodicities of south Florida Littorinidae
(Gastropoda: Prosobranchia).—Bulletin of Marine Science 21(4):826-840.

Clench, W. J., and R. Tucker Abbott. 1942. The genera Tectarius and Echininus in the western
Atlantic.—Johnsonia 1(4):1-4.

Cooke, A. H., A. E. Shipley, and F. R. C. Reed. 1895. Molluscs, Brachiopods (Recent), Brachiopods
(Fossil).—The Cambridge Natural History 3, Macmillan and Co., New York and London,
M+ 7555 pp.

Cotton, B. C. 1947. Some Tertiary fossil mollusks from the Adelaidean Stage (Pliocene) of South
Australia.—Records of the South Australian Museum 8(4):653-670.

Drew, G. A. 1901. The life history of Nucula delphinodonta (Mighels).—Quarterly Journal of Mi-
croscopical Science 44(3) N.S.:313-391.

Fretter, V., and A. Graham. 1962. British prosobranch molluscs.—The Ray Society, London,
wv 75) pp.

Hyman, L. H. 1967. The invertebrates, 6, Mollusca 1. McGraw-Hill, New York, vii + 792 pp.

80 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Jones, G. F. 1963. Brood protection in three Southern California species of the pelecypod Cardita:—
The Wasmann Journal of Biology 21(2):141-148.

Ladd, H. S. 1960. Origin of the Pacific island molluscan fauna.—American Journal of Science,

Bradley Volume 258-A:137—150.

. 1966. Chitons and gastropods (Haliotidae through Adeorbidae) from the Fiesta Pacific

Islands.—Geological Survey Professional Paper 531:1—98, 16 plates.

Mileikovsky, S. A. 1975. Types of larval development in Littorinidae (Gastropoda: Prosobranchia)
of the world oceans and ecological patterns of their distribution.—Marine Biology 30:129-135.

Morton, J. E. 1958. Molluscs. Hutchinson and Co., Ltd., London, 232 pp.

Pfeiffer, Louis. 1839. Bericht uber die Ergebnisse meiner Reise nach Cuba in Winter 1838—1839.—
Archiv fiir Naturgeschichte, Berlin, Funfter Jahrgang 1:346-358.

Philippi, R. A. 1846. Description of a new species of Trochus, and of eighteen new species of
Littorina, in the collection of H. Cuming, Esq.—Proceedings of the Zoological Society of
London for 1845, part XIII: 138-143.

—. 1847-1851. Abbildungen und Beschreibungen neuer oder wenig gekannter Conchylien
3:1-138.

Rosewater, J. 1970. The family Littorinidae in the Indo-Pacific, Part I. The subfamily Littornininae.—

Indo-Pacific Mollusca 2(11):417—506, plates 325-386.

. 1972. The family Littorinidae in the Indo-Pacific, Part II. The subfamilies Tectariinae and

Echinininae.—Indo-Pacific Mollusca 2(12):507—533, plates 388-408.

. 1980. A close look at Littorina radulae.—Bulletin of the American Malacological Union for

1979:5-8, 8 figures.

Roth, A. 1976. Preliminary checklist of the gastropods of Guam.—University of Guam Marine Lab-
oratory Technical Report No. 27, Sea Grant Publication UGSG-76-03:vi + 99.

Thorson, G. 1946. Reproduction and larval development of Danish Marine bottom invertebrates,
with special reference to the planktonic larvae in the Sound (Oresund).—Meddeleser Fra Kom-
missionen For Danmarks Fiskeri—og Havenders@ggelser, serie: Plankton 4(1):1—523.

Turner, R. D. 1966. A survey and illustrated catalogue of the Teredinidae. The Museum of Com-
parative Zoology, Harvard University, vii + 265 pp.

Van der Schalie, H. 1936. Ovoviviparity among mollusks.—The Nautilus 50(1): 16-19.

Wells, F. E., and C. M. Lalli. 1977. Reproduction and brood protection in Caribbean gastropods
Coralliophila abbreviata and C. caribbaea.—Journal of Molluscan Studies 43:79-87.

Yonge, C. M. 1969. Functional morphology and evolution within the Carditacea (Bivalvia).—Pro-
ceedings of the Malacological Society of London 38:493—527.

Department of Invertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
95(1), 1982, pp. 81-88

A NEW DWARF SPHAERODACTYLUS FROM HAITI
(LACERTILIA: GEKKONIDAE)

Richard Thomas

Abstract.—Sphaerodactylus omoglaux is described from the southeastern Cul-
de-Sac Plain and adjacent montane foothills of Haiti. It is a ground dwelling
species of xeric to semi-xeric situations and belongs to a group of Hispaniolan
Sphaerodactylus including S. altavelensis Noble and Hassler, S. armstrongi No-
ble and Hassler, S. cryphius Thomas and Schwartz, S. darlingtoni Shreve, S.
nycteropus Thomas and Schwartz, and S. streptophorus Thomas and Schwartz.

Recent fieldwork in Haiti resulted in the discovery of an undescribed small
species of Sphaerodactylus in the southeastern part of the Cul-de-Sac Plain and
adjacent foothills of the Massif de la Selle. This new sphaerodactyl is related to
the recently described Sphaerodactylus cryphius and S. streptophorus (Thomas
and Schwartz, 1977).

_ Abbreviations.—ASFS refers to the Albert Schwartz Field Series, CM to the
Carnegie Museum of Natural History, RT to Richard Thomas’s personal collec-
tion, UMMZ to the University of Michigan Museum of Zoology, and USNM to
the National Museum of Natural History.

Sphaerodactylus omoglaux, new species
Figs. 1-3

Holotype.—USNM 221840, adult female, taken on the eastern edge of the town
of Fond Parisien near the shore of Etang Saumatre, Dept. de Ouest, Haiti, on
1 August 1979 by Richard Thomas. .

Paratypes.—ASFS V50461-62, 0.7 km E Fond Parisien, 16 July 1978, R. Thom-
as; ASFS V50463, RT 5681, same locality as preceding, 23 July 1978, R. Thomas;
RT 7173, ca. 1 km E Fond Parisien, 2 July 1979, R. Thomas; RT 7682, UMMZ
172100-01, CM 83302, USNM 221841, same data as holotype; RT 5591, 8 km
airline NW Fond Verrettes, ca. 0.3 km by road west of the ford across the Riviere
Soliette, 424 m, 19 July 1978, R. Thomas; all paratypes are from the Dépt. de
Ouest, Haiti.

Diagnosis.—A small species of Sphaerodactylus (Fig. 1), maximum snout-vent
length 20 mm; dorsal scales flattened, keeled, imbricate (30-34 between axilla
and groin), no area of middorsal granules or granular scales; smooth throat and
ventral scales (27-30 between axilla and groin); 51-57 scales around midbody;
dorsal body scales with 5 to 9 hair-bearing scale organs, each with one hair, along
distal edge of scale; rostral with a large flat dorsal area bordered by a rim and
sloping towards the tip; 2 postnasals; large cobble-like snout scales (10-13 across
base of snout). Head (Fig. 2) with a bilobed light anterior figure having postero-
lateral extensions; paired, transverse mid-nuchal marks; paired but separate scap-
ular ocelli; traces of dorsolateral striping; dorsal body pattern of scattered dark

82 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Sih Si ! z
Cr Gk
7
Ale

AL “

AL
Frc ie Foaecgoael a
40 SO ee 25

Fig. 1. Line histograms of midbody scale counts (left side of figure) for Sphaerodactylus omoglaux
(OM), S. cryphius (CR), S. streptophorus (ST, sample from the Pedernales area, Dominican Re-
public), and S. altavelensis (AL). The altavelensis sample above the dashed line is from the area of
sympatry with S. cryphius, that from below the line is from the area of sympatry with S. omoglaux
(Fond Parisien localities). The right side of the figure shows histograms of snout—vent length in mm
size Classes; in all graphs the smallest vertical unit is a single individual.

brown flecks; lineate sacral figure; lineate-ocellate caudal pattern; boldly lined
throat in males; no ventral lines.

Description of holotype.—An adult female, 19 mm snout-vent length, tail (part-
ly regenerated), 13 mm; dorsal body scales flattened, strongly keeled, acute and
imbricate, 31 between axilla and groin; throat scales smooth, not becoming ex-
tremely minute on central throat area; ventral scales smooth, flattened, rounded,
imbricate, 30 between axilla and groin; 52 scales around midbody; lamellae of
fourth toe of left pes 9. Snout moderate, rostral scale with large dorsal flat area
bordered by a rim and sloping toward tip; snout scales broad, subhexagonal,
keeled, cobble-like and subimbricate; | internasal; 2/2 postnasals; 3/3 supralabials
to mid-eye; temporal scales and dorsal head scales keeled, juxtaposed, subim-

VOLUME 95, NUMBER 1 83

Fig. 2. Dorsal patterns of S. omoglaux: A, RT 7686; B, RT 5591; C, USNM 221840 (holotype).

bricate; first infralabial broader anteriorly than posteriorly, subrectangular; dorsal
scales of tail acute, keeled, flattened and flat-lying; ventral scales of tail smooth
and enlarged in median line.

Dorsal ground color in life brown, markings darker brown to nearly black; head
with dark preocular lines and a pale transverse snout bar; pale anterior cephalic
figure bilobed, extending behind eyes onto parietal region and having a short pale
extension from the posterolateral corner of each lobe; a small pale parietal spot
in the space between the posterior parts of the lobes; a pair of pale, dark-edged,
transverse mid-nuchal marks; paired pale (cream) dark-edged scapular ocelli dor-
solaterally positioned and not joined by a scapular patch of dark pigment; broad,
dark-edged dorsolateral stripes present in scapular region, the uppermost stripe
edge on each side interrupted by a scapular ocellus; stripes fading out just beyond
level of axilla; dorsum brown with irregular mottling and spotting of darker
brown; sacral markings of indistinct dark lines; caudal pattern lineate (dorsolat-
eral, lateral, and ventrolateral dark lines) with light spots or ocelli spaced along
the dorsalmost pair; venter unpatterned, pale (off-white) in life.

Variation.—The maximum snout-vent length is 20 mm (female); two egg-bear-
ing females are 19 mm, and the smallest escutcheoned male is 16 mm (largest
male 18 mm). The scale morphology of the paratypes is similar to that of the
holotype. Eight specimens have one internasal; 4 have 2; postnasals are 2/2 in all
but one having 1/2; scales across the snout between the posterior ends of the first

84 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

supralabials are 10 (3 specimens), 11 (4), 12 (5), 13 (1); upper labials to mid-eye
are 3/3 in all; dorsal scales between axilla and groin are 30—33 (¢ = 31.6); ventral
scales axilla to groin, 27— 30 (¥ = 28.8); scales around midbody, 51-56 @ = 53.5);
subdigital lamellae 8 (1), 9 (9), or 10 (2); keeling of throat and ventral scales is
absent in all. The escutcheon of males varies from 5 to 7 scales long and from 15
to 19 scales wide.

In color and pattern the paratypes are generally similar to the holotype. All
have a bilobed anterior head figure, each lobe with a median postpalpebral line
of dark pigment. All but one show some evidence of the posterolateral extensions
of the lobes (in three the extensions are indistinct or irregular). All have the
preocular lines; all but three have the transverse snout bar, either pale as in the
holotype or with the dark posterior edge being the most salient feature and thus
appearing as a dark snout bar; median snout pigmentation varies from pale, nearly
unpigmented, to solidly pigmented with dark brown or pigmented but with a light
center. Nine of the 12 paratypes have plainly evident paired transverse mid-
nuchal marks; in two the marks are obscured by irregular mottling of pigment,
and in one they are absent. All specimens show some indication of the dorsolat-
eral stripes in the scapular area, even if only unilaterally. Paired scapular ocelli
are found in all and are distinctly separate, not interconnected by a dark patch,
although each ocellus is surrounded by a zone of dark pigment (Fig. 2). Dorsal
body coloration varies from almost uniform brown through the presence of iso-
lated small brown flecks on a paler brown ground color to being fairly heavily
mottled with irregular dark scale clusters. The sacral pattern is not prominent
and consists of, when present, the dark dorsal edges of the dorsolateral lines
which reappear in the sacral region. Unregenerated tails show the lineate and
ocellate condition described for the holotype, the most proximal pair of ocelli
often being the most prominent. Throats are heavily streaked with dark brown
lines in the three largest males (the fourth male is very small, and the throat lines
are present but not bold). Females lack throat streaking or show it only very
weakly. Ventral coloration is pale (largely unpigmented) in most, but some have
a peppering of melanophores over much of the surface, concentrated around the
scale edges; the venters are never lined. |

Distribution.—Known only from the southeastern part of the Cul-de-Sac Plain
of Haiti from the region of Fond Parisien southeastward into the foothills of the
La Selle near Soliette (Fig. 3).

Comparisons.—Sphaerodactylus omoglaux belongs to a closely related group
of generally small to medium-sized species that includes darlingtoni Shreve, al-
tavelensis Noble and Hassler, armstrongi Noble and Hassler, streptophorus
Thomas and Schwartz, cryphius Thomas and Schwartz, and nycteropus Thomas
and Schwartz. The last four and S$. omoglaux appear to form a more closely
related cluster within the group (Thomas and Schwartz in press). With the ex-
ception of S. armstrongi, these are small species. These four share polythetically
a suite of characters, and the distributions are largely allopatric or parapatric
(Fig. 3). Although sympatry is likely for at least two of the group, conspecific
relationship between others cannot be ruled out. Because of the commixture of
characters, even in sympatric members of the (larger) group, it is unwise to
assume subspecific relationship without clear evidence of intergradation.

The most pertinent comparisons are with Sphaerodactylus cryphius, known

VOLUME 95, NUMBER 1 85

Fig. 3. Map of a portion of southern Hispaniola. Hexagons indicate S$. omoglaux localities;
rhombs, S. cryphius; circles, S. streptophorus, and triangles, S$. armstrongi.

from the south side of the Valle de Neiba, some 50 km to the southeast, in the
Dominican Republic, and with S. streptophorus of the western, mostly peripheral,
parts of the La Selle-Baoruco massif (Fig. 3). Sphaerodactylus omoglaux seems
to be the ecological equivalent and the geographical vicariant of S. cryphius. In
comparison with omoglaux, cryphius has a reduced color pattern with less elab-
orate and less contrasting markings; the most striking absolute difference between
the two lies in the nature of the scapular ocelli: prominent and separate with no
conjoining patch in omoglaux, absent or small and close-set and with a small,
non-enclosing but conjoining patch in cryphius (Figs. 2, 4). Furthermore the two
taxa are strongly different in midbody scale counts (Fig. 1; Student’s f-test in-
dicates P < 0.01 that the two samples are from the same population). Sphaero-
dactylus cryphius also lacks sexual dichromatism in the throat pattern or any
other feature.

Sphaerodactylus streptophorus is a distinctly larger lizard. Its scale counts
strongly overlap those of omoglaux, but its head pattern differs strongly from
that of omoglaux (Fig. 4A), although it may have a cognate of the anterior ce-
phalic figure (but not bilobed: Fig. 4B). The scapular ocelli, when present, are
separate, as in omoglaux, but not as boldly dark-edged and are sometimes farther
back on the body. The cognate of the mid-nuchal marks of omoglaux in strep-
tophorus is usually a thin transverse light line (whence the specific epithet mean-
ing ‘‘wearing a necklace’’), although it may be broken into two segments. Another
frequent feature of streptophorus is a long diagonal postauricular mark that often
meets part of the cephalic figure to form a mesially directed temporal wedge.

Sphaerodactylus armstrongi is a medium to large-sized member of this group

86 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

/
j
|
|
|
A |

Fig. 4. Semi-diagrammatic anterior dorsal patterns of S. streptophorus (A, ASFS V2580; B, ASFS
V2590) and S. cryphius (C. ASFS V20511).

that occurs in the uplands and easternmost lowlands of the La Selle-Baoruco
ranges. It lacks the bilobed cephalic figure, but it may show a light dorsal head
pattern bounded by postocular stripes and an occipitonuchal M-—shaped figure
(subspecifically variable). The postauricular marks (legs of the M) are probably
homologues of the shorter, posteroventral light marks that extend from the cor-
ners of the cephalic figure in omoglaux. Easternmost S$. armstrongi often has
prominent. separate scapular ocelli somewhat resembling the condition in om-
oglaux. Sphaerodactylus armstrongi further differs in having one postnasal and
a rounded rostral that does not have a sharply delimited flat area and does not
slope towards the tip. Sphaerodactylus armstrongi also differs from omoglaux
(and Streptophorus, cryphius, and nycteropus) in its proportionately larger
ear openings.

Sphaerodactylus nycteropus lacks mid-nuchal marks or scapular ocelli; it does
have, in some specimens, a cephalic figure but no posterolateral marks. Evidence
of dorsolateral lines is present. The venter is strongly lined. In scale characters
and size nycteropus does not differ markedly from omoglaux. Sphaerodactylus
nycteropus is now known from a total of five specimens.

It is pertinent to compare S. omoglaux and S. altavelensis, which occur syn-
topically. Both are small sphaerodactyls of generally similar appearance, although
altavelensis is somewhat larger (Fig. 1). Both have similar scale morphology,
although omoglaux has higher midbody scale counts and lacks the Keeling on the
throat that is present in altavelensis. Reflecting the size difference, the modal
number of subdigital lamellae is 10 in altavelensis, 9 in omoglaux. Both species
have bilobed anterior cephalic figures, and the posterior cephalic figure of alta-
velensis may be the homologue of the mid-nuchal marks of omoglaux. Sphae-
rodactylus altavelensis has small, close-set scapular ocelli that are situated on

VOLUME 95, NUMBER 1 87

the outer edges of an irregular scapular patch. The patch may have a short light
anterior margin giving it a straight border anteriorly and may also have indications
of a larger, enclosing but not delimiting, light border laterally and posteriorly (this
feature and patch size varies in different subspecies of altavelensis; see Thomas
and Schwartz (in press) for a discussion of variation in altavelensis). The bold
throat pattern is lacking in either sex of altavelensis, although faint striping is
present in some individuals, but the venter is lined.

Remarks.—At the two localities near Fond Parisien all of the specimens of
Sphaerodactylus omoglaux were collected in leaf litter and beneath rocks within
about 200 m of the south shore of Etang Saumatre. The habitat is a mixture
of xerophytic scrub (Acacia, Prosopis, Opuntia, Cephalocereus) and more mesic
woods, which include Sabal palms, Swietenia, Mangifera, Tamarindus, Catalpa,
etc. There is a moderate gradient going toward the shore of the lake, where
the habitat becomes more mesic and densely vegetated, although open and closed
situations are patchy due to cutting by man. The only other sphaerodacty] that
we found was S. altavelensis, which occupies the same ground habitat as om-
oglaux. We obtained altavelensis in greater numbers than omoglaux (about 3 to
1). All of the omoglaux were obtained by one person, whereas native collectors
contributed to our sample of altavelensis. Sphaerodactylus altavelensis is com-
mon in all of the leaf-litter habitats in the more or less wooded situations, whereas
omoglaux seems to be confined to shadier, lower canopy (or thicker canopy)
situations. We were unsuccessful in attempting to collect any sphaerodactyl in
the pure desert scrub a few kilometers to the northwest of Fond Parisien, although
some species almost certainly occur there.

Sphaerodactylus omoglaux exists at least macrosympatrically with S. copei,
S. elegans, and §. cinereus, in addition to S. altavelensis, just discussed. The
first three are large species and are inhabitants principally of trees, rock crevices,
or other three-dimensional niches (e.g., wood piles, thatch roofs). The single
specimen of S$. omoglaux from 8 km NW Fond Verrettes was taken from a dead
Agave rosette in xerophytic scrub, where the road runs along a hillside above
the Riviere Soliette at an elevation of 427 m. This locality is about 15 km southeast
of the type-locality and is the most xeric situation in which the species was found.
A specimen that has been referred to S. streptophorus was taken at Soliette,
probably less than a kilometer from the locality for the specimen of omoglaux just
mentioned. The road descends along the xeric hillside, fords the Riviere Soliette,
and then proceeds along a shadier, more mesic riverside area that is Soliette
proper. The surrounding hillsides are open—scrubby or cultivated. The two
species, omoglaux and streptophorus, may coexist in this area, each restricted
to patches of appropriate habitat.

Etymology.—Omoglaux is from the Greek, omos, shoulder, and glaux, owl,
which is derived from glaukos, gleaming, in reference to the eyes. The allusion
is to the prominent, separate scapular ocelli.

Acknowledgments

I wish to thank Luis Rivera Cruz for his capable assistance in the field. The
fieldwork during which the specimens of Sphaerodactylus omoglaux were col-
lected was supported by National Science Foundation Grant SER 77-04629.

88 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Literature Cited

Thomas, R., and A. Schwartz. 1977. Three new species of Sphaerodactylus (Sauria: Gekkonidae)
from Hispaniola.—Annals of the Carnegie Museum of Natural History 46(4):33-43.

, and . In press. Variation in two species of Hispaniolan Sphaerodactylus (Reptilia:
Sauria: Gekkonidae).—Bulletin of the Museum of Comparative Zoology.

Biology Department, University of Puerto Rico, Rio Piedras, Puerto Rico
0093 1.

PROC. BIOL. SOC. WASH.
95(1), 1982, pp. 89-92

AMERIGONISCUS MALHEURENSIS, NEW SPECIES,
FROM A CAVE IN WESTERN OREGON
(CRUSTACEA: ISOPODA: TRICHONISCIDAE)

George A. Schultz

Abstract.—A new species of Amerigoniscus Vandel, a genus with 8 other
species in the United States, is described and illustrated from Malheur Cave in
western Oregon.

The new species is the third trichoniscid and the second in the genus Ameri-
goniscus Vandel (1950) to be described from Oregon. The first species is Ore-
goniscus nearcticus (Arcangeli, 1932) from MacLeary Park, Portland, and the
second is Amerigoniscus rothi (Vandel, 1953a) from 8 mi (12.9 km) east of Gold
Beach, near the California border. Neither was from a cave although both are
blind and have little or no body pigment. The new species described here is from
Malheur Cave in Harney County in western Oregon. It also has no eyes or body
pigment.

The two male specimens were collected by E. H. Gruber and sent to me by
Jerry Lewis, University of Louisville, Kentucky, and I thank him for sending
them to me.

Amerigoniscus Vandel

The genus was recently reviewed by Vandel (1977) who included information
on the four species and added four new ones. This then is the ninth species placed
in the genus. All are from the United States, from Georgia through Texas to
Oregon. Only A. rothi Vandel is not from a cave. Vandel separated the eight
species by using the configurations of the tip of the exopod of male pleopod | of
each species. He illustrated that structure for all eight species. Unfortunately for
the four new species which he described that is all he illustrated, but the four
species are sufficiently different morphologically and geographically so as not to
be confused with the new species described here.

The relation of Amerigoniscus Vandel to Oregoniscus Hatch (1947) as dis-
cussed by Vandel (1953a:177) is still unsettled since O. nearcticus (Arcangeli) is
based on a female. Vandel (1953b) also discussed some of the species of the
genus, and species of related genera of Trichoniscidae. He placed the first de-
scribed species of Amerigoniscus in Caucausonethes Verhoeff and the genus was
included among the primitive members of the family Trichoniscidae in Tribe 1.
However, when later (Vandel 1977) he changed the generic name of the species
from the United States back to Amerigoniscus he did not mention the placement
of the genus in any subgroup within the Trichoniscidae.

Amerigoniscus malheurensis, new species
Figs. 1[A-N, 2A—C

There are no illustrations of the whole animal of any species of the genus.
Unfortunately the two specimens collected here were too distorted to be drawn,
but the appendages are definitive and are illustrated.

90 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Amerigoniscus malheurensis, holotype male 3.5 mm long: A, Detail tip exopod male
pleopod 1; B, Reverse of A; C—G, Pleopods I—5 respectively; H, Peraeopod I (twisted); I, Mandibles;
J, Maxilla 1; K, Maxilliped; L, Fungal spores (small ones same scale as mandibles); M, Antenna 1;
N, Antenna 2.

Description.—Eyeless, pigmentless. Cephalon and body covered with tuber-
clelike scale groups. Few scale groups on top and one row on posterior margin
of cephalon; 2 rows on peraeonal segments, especially apparent on segments I-V;
few on edges of pleonal segments. Antenna 1 with 8 aesthetascs. Antenna 2 short,

VOLUME 95, NUMBER 1 91

A VA Zé,
AK
oe \ FZ Ve
Fee m
Vitter 3 ~~

Fig. 2. Amerigoniscus malheurensis, holotype male: A, Peraeopod VII; B, Uropod; C, Pleotelson.

with 3 flagellar articles. Mandibles each with well developed molar process and
2 setae in setal row. Right mandible with 3 teeth on incisor process; lacinia mobilis
with apical crown of small teeth. Left mandible with 4 teeth on incisor with 2 on
each of 2 sections. Endopod of maxilla 1 with 2 penicillate setae on apex and |
longer subapical compound seta. Exopod with 8 teeth and | broad seta on apex.
Maxilliped with pointed endite and palp of 2 segments with apical segment large,
pointed and fringed with small setae.

Peraeopod I of male with 2 setae on proximal inner margin of propodus (prob-
ably sexually dimorphic character, but females not obtained). Peraeopod VII of
male with 3 large setae on inner margin of propodus; carpus with simple arrange-
ment of setae. Featherlike dactylar organ present on all peraeopods. Pleopod |
with large exopod with configuration of tip unique (Fig. 1A, B). Pleopod 2 with
exopod with medial posterior margin produced; | seta medially placed on pos-
terior margin. Endopod with proximal segment shorter than length of exopod;
distal segment elongate and tapered to truncate end. Pleopods 1|-3 as illustrated.
Uropodal rami each with | large seta at tip. Pleotelson with posterior margin
produced, rounded, and with 4 small setae.

Length.—Both males 3.5 mm long.

Type-locality.—Malheur Cave, about 18 mi (29 km) southeast of New Princeton
(on Oregon route 78), Harney County, Oregon.

Distribution.—Known only from type-locality.

Etymology.—The name malheurensis means from Malheur, the name of the
cave which is the type-locality.

Disposition of types.—The type-specimens have been placed in the National
Museum of Natural History (Smithsonian Institution), holotype male USNM
184664; paratype male USNM 184665.

Discussion.—The species differs from A. rothi in that the configurations of the
tips of the exopods of male pleopods | are quite different (cf. Vandel, 1977:308,
Fig. 1, and Fig. 1A here). Although the various configurations illustrated by
Vandel (1977) are quite different, the best match is between the tip of the exopod
of the new species and that of A. rothi. There are, however, many differences in
other structures if the illustrations of Vandel (1953a) are compared to those in-
cluded here. Vandel illustrated 11 aesthetascs on antenna | and six flagellar ar-
ticles on antenna 2 for A. rothi. There are eight aesthetascs and three flagellar

92 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

articles on the new species. The general shape of the exopods of pleopods 1
(excluding details of tip) have different shapes in the two species. Male pleopods
2 are also differently shaped—in the new species the exopod has the medial
posterior edge or tip produced; it is aright angle in A. rothi. The proximal segment
of the endopod is proportionately shorter and the tip of the distal segment elongate
and narrowed to a truncate, not pointed, end in the new species. There are other
minor differences which can be seen by comparing the drawings of the two
species.

Ecology.—The type-specimen was found under a rotted stick in the flood zone
at the 1325 foot (397.5 m) level. The paratype was found under a plank in the
flood zone at the 1400 foot (420 m) level. The guts from the mouth to the anus
of both specimens were stuffed with fungal spores on which they were apparently
feeding. The spores are two-chambered (Fig. 1L) and most probably from a fun-
gus of the Deuteromycetes, Didymosporae group. No mycelia were present.

Literature Cited

Arcangeli, A. 1932. Isopodi terrestri raccolti dal Prof. Silvestri nel Nord-America.—Bolletino La-
boratoire di Zoologicae Generale e Agraria, Portici 26:121-141.

Hatch, M. H. 1947. The Chelifera and Isopoda of Washington and adjacent regions.—University
of Washington Publications in Biology 10(5):155—274.

Vandel, A. 1950. Campagne spéologique de C. Bolivar et R. Jeannel dans |’ Amérique du Nord (1928).

Isopodes terrestres recueillis par C. Bolivar et R. Jeannel (1928) et le Dr. Henrot (1946).—

Archives Zoologie Expérimentale et Générale 87(3): 183-210.

. 1953a. A new terrestrial isopod from Oregon, Caucasonethes rothi n. sp.—Pacific Science

7(2):175—178.

. 1953b. Remarques systématiques, morphologiques et biogéographiques sur un groupe de

Trichoniscidae Nord-Atlantiques (Crustacés; Isopodes terrestres).—Bulletin du Muséum Na-

tional d’ Histoire Naturelle, Série 2, 25(4):368-375.

. 1977. Les especes appartenant au genre Amerigoniscus Vandel, 1950 (Crustacés, Isopodes,

Oniscoides).—Bulletin de la Societé d’ Histoire Naturelle de Toulouse 113:303-310.

15 Smith St., Hampton, New Jersey 08827.

PROC. BIOL. SOC. WASH.
95(1), 1982, pp. 93-98

A NEW SPECIES OF DEEP-SEA ISOPOD,
STORTH YNGURA MYRIAMAE, FROM THE
WALVIS RIDGE OFF SOUTH AFRICA

Robert Y. George and Noel A. Hinton

Abstract.—A new species of abyssal isopod, Storthyngura myriamae, is de-
scribed. This species was collected at 5220 meters during the French JEAN
CHARCOT expedition to the Walvis Ridge off South Africa. The study also
includes a detailed comparison of the new species with the most closely related
species Storthyngura caribbea from 1260 meters off the Windward Island in the
West Indies.

Introduction

The deep-sea fauna on the western and eastern sides of the Walvis Ridge off
South Africa between latitudes 25° and 35° south was recently investigated. This
study was conducted aboard the R/V Jean Charcot by the French deep-sea group
at the Brest Oceanographic Laboratory under the leadership of Dr. Myriam Si-
buet. The deep-sea samples contained many isopods belonging to more than
fifteen genera including the abyssal genus Storthyngura.

The isopod fauna on both sides of the Walvis Ridge is thus far poorly known.
However, the Walvis Ridge abyssal region is of zoogeographic interest in view
of the theory that the genus Storthyngura_ originated in the Antarctic slope
(George and Menzies 1968b) and the distribution of Storthyngura species 1s some-
what related to the pattern of the Antarctic Bottom Water flow (George, unpub-
lished data). The Walvis Ridge material contained a hitherto undescribed species
of Storthyngura and the new species is named in honor of Myriam Sibuet for
graciously providing the isopod material for study.

Order: Isopoda, Suborder Asellota
Family: Eurycopidae
Storthyngura myriamae, new species
Fig. 1[A—H

Diagnosis.—Storthyngura with cephalon devoid of any spines. First pereonal
somite lacking dorsal spines; somites 2 to 4 having single, prominent, anterior
dorsal spine; in addition, somites 2 to 4 with posterior transverse ridge exhibiting
median tubercle. Somites 5 to 7 displaying a pair of well-developed median dorsal
spines. First pleonal somite lacking spination. Pleotelson with anterior spine and
pair of posterior dorsal tubercles; 2 well-defined lateral spines and triangular apex.
Basis of uropod approximately same length as endopod; exopod about two-thirds
length of endopod.

Material and station data.—Holotype female: length = 21 mm, width 8 mm
(U.S.N.M. Cat. No. 184668).

Type-locality: Angola Basin west of Walvis Ridge, JEAN CHARCOT Station

94

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

N4

,
Q)
dy,
4

ne
Oe

ee
ana? icky
. *s A
: . Cn oe eA VNVIDDDD
f, b
= i ee sf Fi
hai ©

Fig. 1. Storthyngura myriamae: Holotype female, length 21 mm. A, Dorsal view; B, First antenna;
C, Left mandible; D, Maxillule; E, Maxilla; F, Maxilliped; G, Sixth peraeopod; H, Uropod.

DS09, one female specimen, 5220 m, 26°59.9’S—27°00.0'S, 1°06.7’E—1°06.2’E.

Date: 6-Jan-79.

Other localities: JEAN CHARCOT Station CP10, one female specimen, 5210
m, 26°15.9’—27°00.35’'S, 1°07.1’-1°06.4'E. Date: 7-Jan-79 (returned to Dr. Sibuet
for deposition in the Brest Reference Museum); JEAN CHARCOT Station CP12,

VOLUME 95, NUMBER 1 95

2025

-

30°

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oy
era
0
ees
as
too
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oo
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1 \
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Fig. 2. Area map showing the Walvis Ridge separating the Angola and Cape basins. Stations with
S. myriamae shown in solid circles and stations lacking S$. myriamae shown in open circles.

two female specimens, 4660 m, 27°37.6’—27°38.5'S, 0°50.8’—0°51.3’E. Date: 9-Jan-
79. Fragments from Station DSO7 (See Fig. 2).

General Description

Body shape ovate, approximately 3 times longer than wide. Cephalon including
a pair of mounds on dorsal surface. Lateral margin concave; frontal margin trun-
cate.

First pereonal somite without spination. Antero-lateral margin rounded, coxal
plates small and acicular; second pereonal somite having prominent median spine,
antero-lateral margin rounded; coxal plates protuberant and bilobed, anterior lobe
prominent. Posterior ridge on somites 2 to 4 exhibiting median tubercle; somites
3 and 4 each with produced median spine, antero-lateral margins acutely pro-
duced, prominent and bilobed coxal plates with distinct anterior lobes; somites
5, 6, and 7 with paired prominent median spines, concave posterior margins,
lateral margins produced into spine-like process, coxal plates absent.

Pleon with prominent anterior pleonal somite, smooth and lacking any spines;
pleotelson shield-like, with well-developed acute antero-lateral angles and pro-
jecting postero-lateral spines; apex triangular and converging to a point. Pleotel-
son with anterior spine and pair of conical tubercles located anterior to postero-
lateral spines.

First antennae with capacious basal article, lateral margin entire. Peduncle with

96 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

one broad basal article, second article shorter than elongated third article: fourth
article narrow, about one-half length of third article. Flagellum with more than
60 annulate articles. Mandible with prominent palp consisting of 3 articles; second
article slightly longer than first article; third article inflexed into semi-circular
pattern; inner basal margin displaying a row of thick setae, distal margin with 7
to 8 blunt, short spines and terminal setae. Well-developed cutting phase with
pronounced molar and very dense middle row of approximately 20 spines. Incisor
uniform with quadradentate lobe. Maxillule with outer lobe two times as broad
as inner lobe; both lobes with dense setae at distal ends; outer lobe with approx-
imately 15 stout, spine-like setae. Maxilla trilobed with outer lobes slender, each
furnished with 2 to 4 long setae; inner lobe stout with approximately 20 terminal
setae. Maxilliped with 11 coupling hooks on endite; palp with elongate basal
article, somewhat narrower at base; second and third articles expanded, second
article devoid of marginal setae, third article containing approximately 28 mar-
ginal setae on inner margin; fourth article distally produced into lobe one-third
length of article, with about 10 terminal setae; fifth article narrow, with tuft of
apical setae.

Posterior 3 legs morphologically similar; basis elongated, about as long as car-
pus and ischium combined. Merus and propodus expanded, propodus with dense
row of plumose setae on inner and outer margins; dactyl relatively long and
narrow. Uropod with basis as long as endopod; exopod three-fourths length of
endopod.

Morphological Affinities with S. caribbea

In terms of pleotelson configuration, S. myriamae exhibits a superficial resem-
blance to S. unicornalis and in fact, to all member species of the B, Storthyngura
group according to the scheme proposed by George and Menzies (1968b). How-
ever, this species 1S SO Unique in possessing a transverse ridge in somites two to
four which is posterior to the prominent median dorsal spines in all three somites.
This transverse ridge also exhibits a small median tubercle. The only other species
in the genus Storthyngura that has a similar transverse ridge with tubercles on
the second through fourth somites is S. caribbea which was originally described
by Benedict as a species belonging to the genus Eurycope; subsequently,
Richardson (1905) redescribed this species. Wolff (1962) properly affiliated this
species to the genus Storthyngura but treated it as a subspecies of S$. pulchra.
George and Menzies (1968a, b) offered reasons for distinguishing S. caribbea as
a distinct species.

We believe that the new species S. myriamae, found in the Angola Basin at
5220 m, is somehow very closely related to S. caribbea which occurs off the
Windward Island in the West Indies at a depth of 1260 m. We examined the type
specimen from the U.S. National Museum (courtesy of Dr. Tom Bowman), com-
pared the two species, and arrived at the following major morphological differ-
ences between the two species.

Close observation of S. myriamae and §. caribbea revealed a striking similarity
in the shape of the head and general body form. One major difference is found
in the second somite; the antero-lateral angle is produced in S. caribbea but
rounded in $. myriamae. Also, comparison of the mouthparts showed conspic-
uous anatomical differences in the mandible and maxilliped. The palp of the

VOLUME 95, NUMBER 1 97

maxilliped in S. caribbea has a short basal article widened at its base; S. myria-
mae has a palp with an elongated basal article that is narrow at its base.

Both S. caribbea and S. myriamae have single non-lobed coxal plates on the
first somite, but S. caribbea has coxal plates that are far more produced than
those found in S. myriamae. Storthyngura caribbea has elongated dorsal spines
while S. myriamae has spines of a moderate length. There are also differences
between the two species in the shape of the first pleonal somite and the pleotelson.
In S. myriamae the median dorsal length of the first flat pleonal somite is two
times that of its lateral extremity, but in S. caribbea it is of even length through-
out.

We believe that S. myriamae shows a close affinity with S. caribbea, but based
on the morphological differences, S. myriamae is a distinct new species.

Distribution: Angola Basin vs. Cape Basin

The primary purpose of the French deep-sea expedition was to look for any
faunal differences at the abyssal depths between the Angola Basin (on the western
side of the Walvis Ridge) and the Cape Basin (on the eastern side of the Walvis
Ridge, see Fig. 2). Both basins contained oligotrophic clay-like ooze (Sibuet 1980)
but the Angola Basin is rich in CaCO, (70%) and the Cape Basin is poor in CaCO,
(not more than 5%). In terms of bottom currents, the Angola Basin is subject to
possible inflow of the Antarctic Bottom Water whereas the Cape Basin is blocked
by the Walvis Ridge which possibly acts as a topographic and therefore biogeo-
graphic barrier for the expansion of deep-sea fauna from the Weddell Sea—Scotia
Sea Antarctic region. This geographic zone was considered as a center of origin
for the genus Storthyngura (George, 1980). It is also known that the Antarctic
Bottom Water extends north even beyond the equator into the North Atlantic
and abyssal regions off the West Indies.

In this present study we have carefully examined the isopod material from the
deep-sea stations on both sides of the Walvis Ridge, four stations from the Angola
Basin and six stations from Cape Basin (Fig. 2). We encountered Storthyngura
myriamae only in the Angola Basin occurring at all four stations. Storthyungura
myriamae was not found in any one of the six stations at comparable depths in
the Cape Basin. It is of interest to point out that S$. myriamae belongs to Stor-
thyngura B, group which includes six species of similar pleotelson configuration
(S. fragilis from the North Pacific, S. caribbea from the North Atlantic, S. uni-
cornalis from the South Pacific, S. gordonae from the Indian Ocean, and S.
challengeri from the Antarctic Ocean). Although the B, group is not represented
in the Cape Basin, C-group (S. symmetrica) and D-group (S. triplispinosa) are
represented in the Cape Basin (George and Menzies 1968b). Such a pattern of
difference in biogeographic distribution between B-group and C & D-groups sug-
gests apparently different evolutionary pathways for the species within the genus
Storthyngura.

This new species from the Angola Basin shows close morphological similarity
with S. caribbea from the West Indies region which is separated from the Angola
Basin by a series of abyssal basins such as the Guinea Basin, Sierra Leone Basin,
Cape Verde Basin, and Guiana Basin. These basins have a deep sill-depth and
therefore, faunal affiliation between these basins at the abyssal depths should be

98 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

carefully investigated. However, our knowledge of the abyssal zoogeography
and faunistic link between these basins is very limited due to paucity of deep-sea
samples.

It is of interest to point out that S. myriamae is also related to S. challengeri
from the Antarctic Ocean. We can not, however, offer any explanation of the
evolutionary directions and speciation pattern of these closely related B-group
Storthyngura species. On the basis of plesiomorphic features, the Antarctic slope
species from the Weddell Sea region appear to be somewhat primitive (George
1980), although it is difficult to point out which one is the most primitive or
parental species. Nevertheless, the global distribution pattern of Storthyngura
species suggests that their speciation and radiation into the abyssal regions of the
World Oceans since Miocene period is possibly promoted by the flow of the
Antarctic Bottom Water which originates in the Weddell Sea region. This hy-
pothesis obviously calls for careful reevaluation and further investigation.

Literature Cited

George, R. Y. 1980. Antarctica as a center of origin for the deepsea isopod genus Storthyngura.

Symposium on Biology and Evolution of Crustacea, Sydney, Australia (May, 1980).

, and R. J. Menzies. 1968a. Species of Storthyngura (Isopoda) from the Antarctic with de-

scriptions of six new species.—Crustaceana 1!4(3):275-301.

, and . 1968b. Distribution and probable origin of the species in the deep-sea isopod

genus Storthyngura.—Crustaceana 15(2):171—187.

Richardson, H. E. 1905. A monograph on the isopods of North America.—Bulletin of the United
States National Museum 54: 1-727.

Sibuet, M. 1980. A report on Campagne Walvis I, JEAN CHARCOT expedition. December 12, 1978
to January 15, 1979. (Unpublished Report.)

Wolff, T. 1962. The systematics and biology of bathyal and abyssal Isopoda, Asellota.—Galathea
Report 6:7—320.

Institute for Marine Biomedical Research, University of North Carolina at
Wilmington, Wilmington, North Carolina 28403.

PROC. BIOL. SOC. WASH.
95(1), 1982, pp. 99-113

PHYTOPLANKTON DISTRIBUTION ALONG THE EASTERN
COAST OF THE USA IV. SHELF WATERS BETWEEN
CAPE LOOKOUT, NORTH CAROLINA, AND
CAPE CANAVERAL, FLORIDA

Harold G. Marshall

Abstract.—The phytoplankton composition is discussed for southeastern shelf
waters of the United States with the average concentrations at near and far shore
stations given for 328 species. A mixed ultraplankton group, composed of un-
identified species, predominated in numbers, and was part of the shelf assem-
blage of dominant forms with diatoms, dinophyceans, and haptophyceans. Larg-
est cell concentrations were noted near shore with the diatoms and the
ultraplankton component most abundant. Phytoplankton assemblages are given
for near and far shore stations over the shelf.

Phytoplankton composition in the continental shelf waters off the southeastern
coast of the United States has been discussed by Hulburt (1967), Marshall (1969,
1971), and Hulburt and MacKenzie (1971). These studies have indicated diatoms
abundant in the shelf populations with the coccolithophorids and other phytofla-
gellates becoming proportionally more significant beyond the shelf break and in
the Gulf Stream. In addition to diatoms, dinophyceans, and coccolithophores,
representatives from a few other phytoplankton groups have been found common
in these waters. Hulburt and MacKenzie (1971) noted large concentrations of the
cryptophycean Rhodomonas amphioxeia south and north of Cape Hatteras. Dun-
stan and Hosford (1977) reported an abundance of the cyanophyceans Oscillatoria
thiebautii and Oscillatoria erythraea in the south Atlantic bight, with Marshall
(1981) noting 16 cyanophycean species in these shelf waters. Bishop et al. (1980)
indicated little seasonal change in nutrient or phytoplankton concentrations in
this area. They emphasized more variability occurred on the outer shelf which
may be related to intrusions of the Gulf Stream and upwelling along the shelf
break. They also suggested a short-term response time of days to weeks by the
phytoplankton to these events and that responses may differ throughout the year,
resulting in changes in the concentrations among the various populations.

These past studies suggest this section of the continental shelf contains a di-
verse phytoplankton flora with regional variations and changes in population con-
centrations common. The purpose of this study was to evaluate the phytoplankton
composition in this area of approximately 600 km along the shelf between Cape
Lookout, N.C. and Cape Canaveral, Florida. Emphasis was placed on charac-
terizing the populations from near and far shore stations over this section of the
continental shelf.

Methods

Surface water samples were taken from 24 October to 16 November 1973 and
6-20 September 1978 during MARMAP cruises of the South Carolina Marine

100 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Resources Program. These collections were generally taken along transects di-
rected from the coastal area seaward to the vicinity of the shelf break, between
Cape Lookout, N.C. and Cape Canaveral, Florida (Fig. 1). There were 43 stations
in the 1973 collections and 48 stations in 1978. Station depths ranged from 9
m to 318 m, with 45 of the 91 stations located at water depths of less
than 44 m. There were four stations in waters between 200 and 318 m deep.
Reference to near shore stations are those located within 35 km from the
nearest coastline, with those beyond this distance referred to as far shore stations.
In this study 35 of the 91 stations were classified as near shore stations.

At each station a 500 ml surface sample was obtained with a Van Dorn collec-
tion bottle using standard hydrocast procedures. The samples were preserved
immediately with buffered formalin and returned to the laboratory for subsequent
settling. A modified Utermohl method was used with the samples siphoned to a
20 ml concentrate, transferred to a settling chamber and examined with a Zeiss
inverted plankton microscope. The classification format of Hendey (1974), Parke
and Dixon (1976), Drouet and Daily (1956), and Drouet (1968) is mainly followed
in this study. Salinity values and other station data were provided by personnel
from the South Carolina Marine Resources Program.

Results

There were differences in the range and average values for temperature and
salinity for the two cruises (Table 1). The 1973 collections came from water that
was cooler and slightly less saline, with a broader temperature range represented
than what was present in the 1978 samples. The combined average temperature
for both of these cruises was 26.42°C, which was higher than the average fall
temparature (22.4°C) during the previous collections in this area by Marshall
(DTW).

A total of 328 phytoplankters was identified in this study from the two cruises
(Table 2). These consisted of Bacillariophyceae (194), Dinophyceae (83), Hap-
tophyceae (13), Cyanophyceae (16), Euglenophyceae (7), Prasinophyceae (4),
Chlorophyceae (3), Chrysophyceae (5), and Cryptophyceae (3). In addition, there
were high concentrations of an unidentified ultraplankton component that was
divided into three size categories, <3 wm, 3—5 wm, 5-10 wm. These consisted of
round, oval, and irregularly shaped cells, with apparent light green color. Most
numerous was the <3 yum size class which appeared similar to coccoid cyano-
phyceans, whereas the 3—5 wm category resembled chlorophyceans with the 5—10

Table 1.—Temperature and salinity values for surface water at stations during the fall 1973 and
1978 cruises.

Temperature °C Salinity %o
1973 1978 1973 1978
Range 14.9-27.8 27.2—28.6 34.22—-37.07 34.87-38.81
Mean 24.1 27.8 Doel 36.04
Near Shore Mean D3 27.8 35.69 35.84

Far Shore Mean 26.1 27.8 36.03 36.10

VOLUME 95, NUMBER 1 101

g0°
pera = Ve
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Fig. 1. Station locations for water samples along the southeastern continental shelf taken Octo-
ber—November 1973 (®) and September 1978 (@).

102 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Phytoplankton composition and average cell concentrations (no’s/l) for combined stations
in the 1973 and 1978 cruises. Counts for the filamentous cyanophyceans are in filaments per liter.

Stations
Near shore Far shore
BACILLARIOPHYCEAE

Achnanthes longipes Agardh — 0.09
Actinoptychus senarius Ehrenberg 0.29 0.09
Actinoptychus vulgaris Schumann 0.57 0.09
Amphiprora sp. 0.19 —

Amphiprora alata (Ehrenberg) Kutzing — 0.09
Amphora sp. 0.38 —

Amphora binodis Gregory 7.90 0.87
Amphora cuneata Cleve 8.38 —

Amphora egregia var. interrupta Peragallo & Peragallo —_ 0.04
Amphora grevilleana contracta Cleve — 0.17
Amphora marina (W. Smith) Van Heurck 0.38 1.04
Amphora obtusa Gregory 0.19 0.09
Amphora ostrearia Brebisson 4.00 O52
Amphora ovalis Kutzing 0.95 0.70
Amphora peragalli Cleve — 0.17
Amphora terroris Ehrenberg 4.57 —

Asterionella glacialis Castracane 3.05 2.09
Asterionella kariana Grunow — 4.00
Asteromphalus flabellatus (Brebisson) Greville 2.10 0.43
Auliscus sculptus (W. Smith) Ralfs — 0.09
Bacteriastrum comosum Pavillard — 1.13
Bacteriastrum delicatulum Cleve 4.38 9.74
Bacteriastrum hyalinum Lauder 3) {02 12.91
Bacteriastrum hyalinum var. princeps (Castracane) Ikari — 0.17
Bacteriastrum varians Lauder — 0.96
Biddulphia alternans (Bailey) Van Heurck 97.05 0.57
Biddulphia aurita (Lyngbye) Brebisson 4.67 0.17
Biddulphia longicruris Greville 0.57 0.52
Biddulphia mobiliensis (Bailey) Grunow ZOD 0.52
Biddulphia sinensis Greville 0.10 0.04
Biddulphia tridens (Ehrenberg) Ehrenberg 0.76 —

Campylodiscus limbatus Brebisson — 0.09
Campylodiscus rutilis Skvortzow — 0.04
Chaetoceros affine Lauder WoQZ 1.0

Chaetoceros atlanticum Cleve 6.48 6.26
Chaetoceros breve Schutt 18) 28.48
Chaetoceros coarctatum Lauder 10.86 3.45
Chaetoceros compressum Lauder — 0.70
Chaetoceros convolutum Castracane — 0.52
Chaetoceros curvisetum Cleve = 0.35
Chaetoceros decipiens Cleve 63.24 25.87
Chaetoceros densum Cleve 0.38 es
Chaetoceros didymum Ehrenberg — 0.09
Chaetoceros diversum Cleve DD) 3.48
Chaetoceros gracile Schutt — 0.35
Chaetoceros laciniosum Schutt — 0.26
Chaetoceros lorenzianum Grunow 22.86 P17)
Chaetoceros messanense Castracane 0.57 —

Chaetoceros pelagicum Cleve 11.43 —

Chaetoceros pendulum Karsten — 1.04

VOLUME 95, NUMBER 1

Table 2.—Continued.

Chaetoceros peruvianum Brightwell
Chaetoceros pseudocurvisetum Mangin
Chaetoceros radians Schutt

Chaetoceros sociale Lauder

Chaetoceros wighami Brightwell
Climacodium frauenfeldianum Grunow
Cocconeis sp.

Cocconeis molesta var. crucifera Grunow
Cocconeis pinnata Gregory

Corethron criophilum Castracane
Coscinodiscus argus Ehrenberg
Coscinodiscus asteromphalus Ehrenberg
Coscinodiscus centralis Ehrenberg
Coscinodiscus gigas Ehrenberg
Coscinodiscus grani Gough
Coscinodiscus granulosus Grunow
Coscinodiscus lineatus Ehrenberg
Coscinodiscus marginatus Ehrenberg
Coscinodiscus nitidus Gregory
Coscinodiscus nobilis Grunow

~ Coscinodiscus perforatus Ehrenberg
Coscinodiscus radiatus Ehrenberg
Coscinodiscus stellaris var. symbolophora (Grunow) Jorgensen
Coscinodiscus wailesii Gran and Angst
Cyclotella sp.

Cyclotella meneghiniana Kutzing
Cylindrotheca closterium (Ehrenberg) Reiman and Lewin
Cymatosira belgica Grunow

Cymatosira lorenziana Grunow

Dactyliosolen antarcticus Castracane
Dactyliosolen mediterraneus Peragallo

Diploneis crabro Ehrenberg

Diploneis crabro var. pandura (Brebisson) Cleve
Ditylum brightwellii (West) Grunow

Eucampia zoodiacus Ehrenberg

Eunotia sp.

Eunotia bidentula W. Smith

Fragilaria sp.

Fragilaria crotonensis Kitton
Fragilariopsis cylindrus (Grunow) Helmcke and Krieger
Grammatophora sp.

Grammatophora angulosa Ehrenberg
Grammatophora marina (Lyngbye) Kutzing
Guinardia flaccida (Castracane) Peragallo
Gyrosigma sp.

Gyrosigma balticum (Ehrenberg) Cleve
Hantzschia marina (Donkin) Grunow
Hemiaulus hauckii Grunow

Hemiaulus membranaceus Cleve
Hemiaulus sinensis Greville

Stations

Near shore

6.10
2.86

103

Far shore

6.39
0.43
1.65
0.35
UBigeh|\

104 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Continued.

Stations
Near shore Far shore

Leptocylindrus danicus Cleve 19.24 WA
Licmophora sp. 0.19 0.39
Licmophora flabellata (Carmichael) Agardh 3.81 0.09
Mastogloia smithii Thwaites 10.29 0.48
Melosira distans (Ehrenberg) Kutzing 49.81 29.57
Melosira granulata (Ehrenberg) Ralfs 17.33 12.09
Melosira granulata var. angustissima Muller 22.14 85.74
Melosira islandica Muller — 4.75
Melosira moniliformis (Muller) Agardh = 0.04
Melosira nummuloides (Dillwyn) Agardh — 57
Navicula sp. 0.38 0.13
Navicula abrupta (Gregory) Cleve — 0.26
Navicula annulata Grunow 55.05 6.83
Navicula cancellata Donkin IOs57/ 1.74
Navicula clavata Gregory 16.19 1.65
Navicula forcipata Greville 25-33 0.52
Navicula lyra Ehrenberg 7k 0.70
Navicula opima (Grunow) Cleve 0.38 6.35
Navicula praetexta Ehrenberg 0.09 —

Navicula pusilla W. Smith — 0.17
Nitzschia sp. — 0.35
Nitzschia angularis W. Smith 4.19 0.43
Nitzschia distans Gregory 2, 0.61
Nitzschia insignis Gregory ESP —

Nitzschia longissima (Brebisson) Ralfs 16.19 8.00
Nitzschia lorenziana var. densistriata (Peragallo and Peragallo) Hustedt 1.14 1.39
Nitzschia lorenziana var. incerta Grunow 0.76 0.78
Nitzschia lorenziana var. subtilis Grunow 0.76 —

Nitzschia panduriformis Gregory 0.76 0.43
Nitzschia pungens Grunow 56.19 2.96
Nitzschia seriata Cleve —_— 6.78
Nitzschia sigma (Kutzing) W. Smith 0.38 ; —

Nitzschia sigma var. intercedens Grunow — 0.87
Nitzschia socialis Ralfs — 0.17
Nitzschia spathulata Brebisson 4.29 —

Paralia sulcata (Ehrenberg) Cleve 531.05 87.09
Pinnularia sp. — 0.26
Plagiogramma staurophorum (Gregory) Heilberg 448.00 15s.) I)
Plagiogramma vanheurckii Grunow — 0.70
Planktoniella sol (Wallich) Schutt — 0.43
Pleurosigma sp. 0.19 0.52
Pleurosigma angulatum (Quekett) W. Smith 5.24 2.09
Pleurosigma elongatum W. Smith 3.05 1.09
Pleurosigma hamuliferum Brun 30.29 4.51
Pleurosigma nicobaricum (Grunow) Grunow od ll —

Pleurosigma normanii Ralfs 3.43 —

Pleurosigma obscurum W. Smith 152 —

Podosira stelliger (Bailey) Mann 0.19 —

Rhaphoneis amphiceros Ehrenberg 3.43 1522
Rhaphoneis surirella (Ehrenberg) Grunow 9.14 Balls)

Rhizosolenia acuminata (Peragallo) Gran 229 0.26

VOLUME 95, NUMBER 1

Table 2.—Continued.

Stations
Near shore Far shore
Rhizosolenia alata Brightwell TAS SD 336.46
Rhizosolenia alata f. gracillima (Cleve) Grunow 99.62 19.65
Rhizosolenia alata f. indica (Peragallo) Gran 1029.62 404.04
Rhizosolenia bergonii Peragallo 1F33 0.52
Rhizosolenia calcar-avis Schultze Bi-Lo2 31.65
Rhizosolenia castracanei Peragallo 0.57 DD,
Rhizosolenia delicatula Cleve 0.76 7.26
Rhizosolenia fragilissima Bergon 0.57 3750
Rhizosolenia hebetata f. hiemalis Gran 0.38 0.09
Rhizosolenia hebetata f. semispina (Hensen) Gran — AGI
Rhizosolenia imbricata Brightwell 53553 23.28
Rhizosolenia robusta Norman Baill ao0)
Rhizosolenia setigera Brightwell 84.94 14.61
Rhizosolenia stolterfothii Peragallo 37.81 167.78
Rhizosolenia styliformis Brightwell 25.24 11.96
Rhizosolenia temperei Peragallo 0.10 1.83
Schroederella delicatula (Peragallo) Pavillard —- 0.78
Skeletonema costatum (Greville) Cleve 36.10 0.96
Stephanopyxis palmeriana (Greville) Grunow 3.43 0.43
Stephanopyxis turris (Greville) Ralfs SAS 8.04
Surirella crumena Brebisson — 0.09
Surirella pandura var. contracta Peragallo and Peragallo 0.19 —
Synedra crystallina (Agardh) Kutzing 1.90 1.04
Synedra fulgens (Greville) W. Smith 70.10 27.48
Synedra gaillonii (Bory) Ehrenberg 0.95 —
Synedra robusta Ralfs 0.38 —
Synedra tabulata (Agardh) Kutzing — 0.09
Synedra toxoneides Castracane 0.19 0.09
Synedra undulata Bailey IS3e52 1.00
Tabellaria fenestrata var. asterionelloides Grunow 97.52 2522.
Thalassionema nitzschioides Hustedt 808.67 144.70
Thalassiosira baltica (Grunow) Ostenfeld — 0.43
Thalassiosira eccentrica (Ehrenberg) Cleve 13.62 2.09
Thalassiosira gravida Cleve 4.38 1.74
Thalassiosira nordenskioeldii Cleve 25a 1.61
Thalassiosira subtilis (Ostenfeld) Gran — 0.13
Thalassiothrix delicatula Cupp — 1.30
Thalassiothrix frauenfeldii Grunow 99.43 19.61
Thalassiothrix mediterranea Pavillard 12.00 6.63
~Triceratium favus Ehrenberg 0.10 0.13
Triceratium formosum var. pentagonalis (Schmidt) Hustedt 0.19 —
Tropidoneis lepidoptera (Gregory) Cleve 0.19 0.04
Tropidoneis seriata Cleve 0.57 0.35
Unidentified diatoms 201.14 54.43
DINOPHYCEAE
Amphidinium acutissimum Schiller 0.38 8.78
Amphidinium acutum Lachmann — 0.04
Amphidinium bipes Herdman — 0.91
Amphidinium globosum Schroder — 0.52

Amphidinium klebsii Kofoid and Swezy — 0.39

106 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Continued.

Stations
Near shore _ Far shore

Amphidinium lanceolatum Schroder — 0.04
Amphidinium schroederi Schiller — 3.04
Ceratium contortum (Gourret) Cleve 0.19 0.04
Ceratium contortum var. karsteni (Pavillard) Sournia 0.19 0.39
Ceratium digitatum Schutt — 0.04
Ceratium extensum (Gourret) Cleve 0.19 0.48
Ceratium furca (Ehrenberg) Claparede and Lachmann 11.90 3.83
Ceratium fusus (Ehrenberg) DuJardin TAM 0.93
Ceratium geniculatum (Lemmermann) Cleve 0.38 ==
Ceratium horridum (Cleve) Gran 0.76 —
Ceratium kofoidi Jorgensen — 0.09
Ceratium lineatum (Ehrenberg) Cleve 4.76 3.30
Ceratium longirostrum Gourret 0.19 —
Ceratium macroceros (Ehrenberg) VanHoffen 0.38 —
Ceratium massiliense (Gourret) Jorgensen 3.33 0.83
Ceratium minutum Jorgensen — 0.09
Ceratium pentagonum Gourret 0.38 1.83
Ceratium ranipes Cleve — 0.09
Ceratium setaceum Jorgensen — 0.17
Ceratium teres Kofoid — 0.22
Ceratium trichoceros (Ehrenberg) Kofoid 2.48 3.43
Ceratium tripos (Muller) Nitzsch 0.48 0.39
Ceratium tripos var. atlanticum (Ostenfeld) Paulsen 0.57 0.78
Dinophysis caudata Kent Ue DD,
Glenodinium sp. — 0.26
Gonyaulax diegensis Kofoid — 0.43
Gonyaulax fragilis (Schutt) Kofoid = 0.09
Gonyaulax minuta Kofoid and Michener 0.76 0.09
Gonyaulax monilata Howell 29.14 —
Gymnodinium sp. #1 — 0.70
Gymnodinium sp. #2 — 0.35
Gymnodinium coeruleum Dogiel — 0.09
Gymnodinium danicans Campbell 0.76 . Vist
Gymnodinium variabile Herdman 1:52 0.57
Gyrodinium sp. 1.7] 0.09
Gyrodinium dominans Hulburt == 0.17
Gyrodinium estuariale Hulburt — ee
Gyrodinium fusiforme Kofoid and Swezy 36.57 0.70
Heterocapsa triquetra (Ehrenberg) Stein — ety!
Katodinium sp. — 0.70 ©
Katodinium asymetricum (Massart) Fott — 0.09
Katodinium rotundatum (Lohmann) Loeblich IS. 12.85
Oxytoxum gracile Gran a 0.61
Oxytoxum sceptrum (Stein) Schroder 0.19 —
Oxytoxum variabile Schiller = 0.22
Peridiniopsis assymetrica Mangin — 0.09
Podolampas bipes Stein — 0.61
Podolampas elegans Schutt 0.19 —
Podolampas palmipes Stein — 0.09
Podolampus curvatus Schiller — 0.09

Prorocentrum aporum (Schiller) Dodge 17.90 0.52

VOLUME 95, NUMBER 1

Table 2.—Continued.

Stations
Near shore Far shore

Prorocentrum balticum (Lohmann) Loeblich 0.76 0.22
Prorocentrum cassubicum (Woloszynska) Dodge — 0.09
Prorocentrum compressum (Bailey) Abe 9.43 1.04
Prorocentrum gracile Schutt — 0.17
Prorocentrum lima (Ehrenberg) Dodge 0.38 —
Prorocentrum maximum (Gourret) Schiller 0.76 —
Prorocentrum micans Ehrenberg 168.48 9.09
Prorocentrum nanum Schiller 6.38 3.83
Protoperidinium sp. — 0.09
Protoperidinium biconicum (Dangeard) Balech — 0.04
Protoperidinium breve (Paulsen) Balech 0.19 0.13
Protoperidinium brochii (Kofoid and Swezy) Balech 0.10 0.09
Protoperidinium cerasus (Paulsen) Balech 1.14 0.09
Protoperidinium claudicans (Paulsen) Balech 0.19 0.09
Protoperidinium conicum (Gran) Balech Mes) 0.26
Protoperidinium crassipes Kofoid — 0.35
Protoperidinium depressum (Bailey) Balech 5.14 0.78
Protoperidinium divergens (Ehrenberg) Balech 0.76 —
Protoperidinium grande (Kofoid) Balech — 0.04
Protoperidinium leonis (Pavillard) Balech 0.38 =
‘Protoperidinium oceanicum (VanHoffen) Balech — (5345)
Protoperidinium pendunculatum (Schutt) Balech 0.57 0.04
Protoperidinium pentagonum (Gran) Balech 0.57 0.09
Protoperidinium quarnerense (Schroder) Balech — 0.17
Protoperidinium solidicorne (Mangin) Balech — 0.04
Protoperidinium sphaericum (Okamura) Balech 0.38 0.43
Protoperidinium steinii (Jorgensen) Balech — 0.96

Unidentified dinoflagellate cysts 0.19 0.70

Unidentified dinoflagellates 62.10 49.87

HAPTOPHYCEAE

Acanthoica aculeata Kamptner 0.76 0.43
Chrysochromulina sp. — 0.09
Cyclococcolithus leptoporus (Murray and Blackman) Kamptner 14.29 28s ou
Emiliania huxleyi (Lohmann) Hay and Mohler 50.86 60.48
Gephyrocapsa oceanica Kamptner — 12.96
Hymenomonas carterae (Braarud and Fagerland) Braarud 0.38 0.09
Rhabdosphaera claviger Murray and Blackman 3.05 1.04
Rhabdosphaera hispida Lohmann 3.43 _
Rhabdosphaera stylifer Lohmann 15.62 —
Syracosphaera sp. 0.19 —
Syracosphaera molischii Schiller — 0.09
Syracosphaera pirus Halldal and Markali 0.76 0.26
Syracosphaera pulchra Lohmann 1.90 2.70

Unidentified coccolithophorids 469.90 127.48

CHRYSOPHYCEAE

Calycomonas ovalis Wulff 0.09 —
Dictyocha fibula Ehrenberg 50.14 4.00
Distephanus speculum (Ehrenberg) Haekel 0.38 0.35
Ochromonas sp. 158.29 ee

Ochromonas caroliniana Campbell 0.19 —

108 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Continued.

Stations
Near shore . Far shore
CYANOPHYCEAE
Agmenellum quadruplicatum (Meneghini) Brebisson 12.38 —
Agmenellum thermale (Kutzing) Drouet and Daily 4.19 —
Anacystis aeruginosa Drouet and Daily 38.48 —
Anacystis dimidiata (Kutzing) Drouet and Daily 87.62 12.43
Anacystis marina (Hansg) Drouet and Daily 543.05 29.83
Entophysalis deusta (Meneghini) Drouet and Daily DDS) —
Gomphosphaeria aponina Kutzing 45.81 21.43
Johannesbaptistia pellucida (Dickie) Taylor and Drouet 395.85 35.83
Nostoc commune Vaucher 500.95 126.67
Oscillatoria sp. 0.76 —
Oscillatoria erythraea (Ehrenberg) Kutzing 119.90 116.87
Oscillatoria submembranacea Ardissone and Strafforello NS 1.74
Richelia intracellularis Schmidt — 3.13
Schizothrix calcicola (Agardh) Gomont he 52 —
Schizothrix tenerrima (Domont) Drouet 2229 —
Spirulina subsalsa Oersted 0.57 —
EUGLENOPHYCEAE
Euglena sp. 0.19 —
Euglena ehrenbergii Klebs 0.19 —
Euglena fusca (Klebs) Lemmermann 0.19 22
Eutreptia lanowii Steuer 1.71 0.35
Eutreptia viridis Perty 0.95 0.17
Trachelomonas sp. 0.10 0.09
Trachelomonas hispida (Perty) Stein 0.38 0.17
CHLOROPHYCEAE
Chlorella sp. 8.57 1.04
Crucigenia tetrapedia (Kirchner) West and West 1.90 0.87
Staurastrum quadricuspidatum Turner 0.76 —
PRASINOPHYCEAE
Pyramimonas sp. 0.19 —
Pyramimonas torta Conrad and Kuff — 0.09
Tetraselmis sp. 0.67 —
Tetraselmis gracilis (Kylin) Butcher — 0.09
CRYPTOPHYCEAE
Chroomonas sp. 0.09 —
Cryptomonas sp. 0.38 3.39
Cryptomonas sp. #2 0.09 —
OTHERS
Unidentified green cells (<3.0 microns) 32,154.76 12,844.57
Unidentified green cells (3—5 microns) 105.95 290.22

Unidentified green cells (S—10 microns) 3.05 2.61

VOLUME 95, NUMBER 1 109

Table 3.—Average concentrations of cells (no’s/l) for the various phytoplankton groups at near and
far shore stations in 1973 and 1978.

Near shore Far shore

1973 1978 1973 1978 Combined
Bacillariophyceae 7850 4753 1656 2073 3161
Dinophyceae 621 83 160 33 185
Haptophyceae 949 113 78 126 Dy,
Chrysophyceae 89 <] 10 <|| 7)
Cyanophyceae Zs 1037 396 299 778
Euglenophyceae 8 1 <i <I p)
Chlorophyceae <1 iy <1 D) 3
Cryptophyceae <il <| 3 3 2D
Prasinophyceae 1 Kil <| <I <\|
Ultraplankton* 4952 56,642 4158 20,435 es 22

* Combined size groups of unidentified cells less than 10 microns in size.

jum group more indiscriminate to a specific taxonomic group. In addition to these
non-flagellated forms, there were also some phytoflagellates included in these size
categories. An ultraplankton component to estuarine and marine habitats has
been recognized in recent years (Malone 1971; McCarthy et al. 1974; among
others). These cells may include the cyanobacteria (blue-green algae) found
widely distributed in the western north Atlantic by Waterbury et al. (1979), and
Johnson and Sieburth (1979). They are also similar to those found in the Chesa-
peake Bay plume by Marshall (in press) and off the northeastern U.S. coast by
Marshall and Cohn (1981).

The study area represents a broad, crescent-shaped segment of the southeast-
ern continental shelf, that reaches its greatest width in the area between Jack-
sonville and Savannah. In this region the shelf break is approximately 120 km
from the coast. The phytoplankton populations varied over the shelf with distinct
groups more characteristic of either the near or far shore stations. Average total
concentration of cells was generally greater near shore as was the presence of
the taxonomic groups represented in the samples, with the exception of the cryp-
tophyceans, haptophyceans, and an unidentified ultraplankton component (Table
3). During both cruises, the far shore populations of the cryptophyceans, although

Table 4.—Numbers and percentages of species within each group that were noted limited to near
and far shore stations, or found at both stations.

Only at Only at

Total near shore far shore In both areas
Bacillariophyceae 194 "32 17% 53 27% 109 56%
Dinophyceae 83 12 14% 39 46% 32 40%
Haptophyceae 13 3 23% 4 31% 6 46%
Chrysophyceae 5 2 40% 0 3 60%
Cyanophyceae 16 8 50% 1 6% 7 44%
Euglenophyceae dL 2 29% 0 5) 71%
Chlorophyceae 3 1 33% 0 2 67%
Cryptophyceae 3 2 67% 0 1 33%
Prasinophyceae 4 2 50% 2 50% 0

110 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

not high, were greater than what was found at near shore stations. However, the
total phytoplankton composition was dominated by diatoms, dinoflagellates, hap-
tophyceans, cyanophyceans, and the ultraplankton component during both cruis-
es over the shelf. The total near shore populations included more of the smaller
sized diatoms, whereas over the central and far shelf, larger sized diatoms were
abundant. Of the 194 diatoms, 56% of the species were found at both near and
far shore stations, with another 17% limited to the near shore and 27% noted only
at far shore stations (Table 4). The prominent diatoms over the shelf were Cy-
matosira belgica, Paralia sulcata, Plagiogramma staurophorum, Rhizosolenia
alata, R. alata indica, R. stolterfothii, and Thalassionema nitzschioides (Table 5).
Rhizosolenia alata averaged 713 and 404 cells/l at near and far shore stations,
with R. alata indica averaging 1029 and 404 cells/l at near and far shore sites. In
1978, Rhizosolenia alata indica reached 18,472 cells/l directly off Savannah, with
other “‘pockets’’ of high concentration scattered over the shelf. These two
species, with Rhizosolenia alata gracillima, R. calcar avis, R. hebetata semis-
pina, R. setigera, R. stolterfothii, and R. styliformis represented a common dia-
tom and generic assemblage throughout the shelf.

The phytoflagellates were not found in high concentrations in these collections
but were generally widely distributed. Only 14% of the dinophyceans (Pyrrho-
phyceans) were limited to the near shore stations, with 40% of the species found
at both near and far shore sites and 46% limited to the far shore stations. The
haptophyceans, (Prymnesiophyceans) consisted mostly of coccolithophores, with
46% of this class common across the shelf and 31% of the species limited to the
far shore sites. The euglenophyceans, chlorophyceans, and chrysophyceans were
found in low concentrations, but widely distributed over the shelf, with cyano-
phyceans, and prasinophyceans more common near shore. The prominent near
shore dinophyceans were Prorocentrum micans, P. aporum, Gyrodinium fusi-
forme, Gonyaulax monilta, with Ceratium furca, Dinophysis caudata, and Ka-
tyodinium rotundatum common over the entire shelf. Other characteristic near
shore species include Emiliania huxleyi, Ochromonas sp., Dictyocha fibula, An-
acystis marina, and Johannesbaptistia pellucida. A far shore dominant was Emi-
liania huxleyi, with Cyclococcolithus leptoporus and Oscillatoria erythraea found
over the shelf in significant numbers. There was no increase in cell concentrations
at stations near the shelf break during either of these cruises. Total phytoplankton
populations were generally low, but gave evidence of patchiness in cell concen-
trations over the entire area during both cruises.

Discussion

The shelf phytoplankton possessed a diverse assemblage of 328 species from
ten taxonomic categories. Diatoms, dinophyceans, haptophyceans, cyanophy-
ceans, and the ultraplankton component represented the most abundant forms
and were distributed over the entire shelf. Each of these categories had charac-
teristic species in the near and far shore stations, with several species common
to both (e.g. Rhizosolenia alata, R. alata indica, Emiliania huxleyi, etc.). The
cyanophyceans were more concentrated in the near shore areas, with many
species widely distributed over the shelf. It is also suggested that many of the
ultraplankters (size <3 wm) are cyanophyceans which may indicate added sig-

VOLUME 95, NUMBER I

111

Table 5.—Prominent phytoplankton associated with near and far shore stations during the 1973 and

1978 collections.
Near shore assemblage

Diatoms

Biddulphia alternans
Cymatosira belgica
Chaetoceros decipiens
Fragilariopsis cylindrus
Cylindrotheca closterium
Guinardia flaccida
Hemiaulus sinensis

Melosira distans

Paralia sulcata
Plagiogramma staurophorum
Rhizosolenia alata
Rhizosolenia alata gracillima
Rhizosolenia alata indica
Rhizosolenia setigera
Synedra fulgens

Synedra undulata

Tabellaria fenestrata asterionelloides
Thalassionema nitzschioides

Far shore assemblage

Diatoms

Chaetoceros decipiens
Climacodium frauenfeldium
Guinardia flaccida

Rhizosolenia alata

Rhizosolenia alata indica
Rhizosolenia stolterfothii
Thalassionema nitzschioides
Melosira granulata angustissima

Dinophyceans

Ceratium furca
Dinophysis caudata
Katyodinium rotundatum

Others

Emiliania huxleyi
Cyclococcolithus leptoporus
Oscillatoria erythraea
Cryptomonas sp.

Thalassiosira frauenfeldii Ultraplankton component

Dinophyceans
Gonyaulax monilta
Gyrodinium fusiforme
Prorocentrum micans
Prorocentrum aporum
Ceratium furca
Dinophysis caudata
Katodinium rotundatum

Others

Emiliania huxleyi
Ochromonas sp.

Dictyocha fibula

Anacystis marina
Johannesbaptistia pellucida
Ultraplankton component

nificance in the presence of this group to the region. These cells also appear
widely distributed over the eastern shelf of the United States (Marshall and Cohn,
1981). Other representatives within this size category and the 3-5 um group
appear to include chlorophyceans, among other types. Many of these phyto-
plankters may represent the so called ‘‘lesser’’ systematic categories (rather than
the dinophyceans, diatoms, and haptophyceans). Of note are the cryptophyceans,
whose concentrations were not high in these collections in comparison to the
more prominant groups, but maintained a broad distribution pattern over the
Shelf. However, the total significance of this and other groups is not completely
represented in this study. Although the present collections include an extensive
shelf coverage from two different cruises, there was a temporal limitation of the

112 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

data (both cruises were in the fall and each covered a limited time span for the
collections). Some smaller scale variations in composition and concentrations
also occurred along the transects between adjacent stations. This patchiness in-
volved high concentrations of one or more species at scattered locations during
both cruises and was more typically found among species of diatoms and dino-
flagellates. There was also a pattern of high cell counts for the ultraplankton
component at stations nearest the estuaries with decreasing concentrations sea-
ward. However, the contributions of this unidentified ultraplankton component
to the biomass of the standing crop appeared to be consistently small in com-
parison to the amount attributed by diatoms and dinoflagellates. Unknown is the
long term impact of these cells to the productivity and annual biomass totals for
this area.

Pollen and moth scales were also common in the samples and both were noted
at stations up to 100 km off the coast. Rhaphoneis surinella was found only
attached to grains of sediment that were collected in the water column. Richelia
intracellularis was not found in a free state but as an endosymbiont in Rhizoso-
lenia styliformis and R. hebetata semispina. Epibiotic fungi (chytridiaceans) were
also noted on Rhizosolenia alata indica. The reported dinoflagellate cysts con-
sisted mainly of Protoperidinium pentagonum.

In summary, the results from collections at 91 shelf stations indicated higher
concentrations for each taxonomic group, except the cryptophyceae, and hap-
tophyceans, near shore during both collection periods. This pattern was also
noted for the unidentified ultraplankton component. Exceptions to this pattern
were common, and occurred as patchiness of species dominance and major shifts
in individual concentration values along transects. The shelf populations were
dominated by numerous large sized centric diatoms (e.g. Rhizosolenia alata and
R. alata indica), with clusters and single filaments of the cyanophycean Oscil-
latoria erythraea also common. However, numerous small sized cyanophyceans,
diatoms and phytoflagellates were also characteristic of the shelf region. Another
prominent group was the cyanophyceans, these were widespread over the shelf,
often illustrating patchiness and high concentrations of cells near estuaries. Mar-
shall (1981) associates many of the coastal marshes bordering the inner shelf area
as a possible origin for many of these cyanophyte species.

Not observed in this study was the pattern of increased cell concentrations
from the middle to outer shelf areas, as noted by Bishop ef al. (1980). However,
a similar increase in cell concentrations near the shelf break was noted by Mar-
shall (in press) off the Virginia coast. In contrast, the results at shelf stations
from these cruises were generally uniform in their composition and concentra-
tions, with occasional patchiness found broadly distributed, but more centered
near and slightly beyond coastal estuaries. However, it is noted that these cruises
were both limited to fall collections and covered a brief collection period.

Acknowledgments

Appreciation is given to the National Marine Fisheries Service (MARMAP
Program) in cooperation with the South Carolina Marine Resources Institute of
Charleston, South Carolina, which provided the samples and supplementary station
data, with special thanks to Drs. Victor Burrell and Charles Baran. Further ap-

VOLUME 95, NUMBER 1 113

preciation is given to graduate assistants Charles K. Rutledge, Stephen Cibik,
Brad Fawley, and Laurie Kalenak for their contributions in this study, and to Dr.
Paul Kirk for identification of the chytridiacean fungi.

Literature Cited

Bishop, S., J. Yoder, and G. Paffenhofer. 1980. Phytoplankton and nutrient variability along a cross-
shelf transect of Savannah, Georgia, U.S.A.—Estuarine and Coastal Marine Science
2:359-368.

Drouet, F. 1968. Revision of the classification of the Oscillatoriacea. Monogr. 15.—The Academy of

Natural Sciences of Philadelphia, Fulton Press, Lancaster, Pa., 370 pp.

, and W. A. Daily. 1956. Revision of the coccoid Myxophyceae.—Butler University Botanical

Studies. Vol. XII. Hafner Press, New York, N.Y., 222 pp.

Dunstan, W., and J. Hosford. 1977. The distribution of planktonic blue green algae related to the
hydrography of the Georgia Bight.—Bulletin of Marine Science 27:824-829.

Hendey, N. I. 1974. A revised check-list of British marine diatoms.— Journal of the Marine Biological
Association United Kingdom 54:277-300.

Hulburt, E. M. 1967. Some notes on the phytoplankton off the southeastern coast of the United

States.—Bulletin of Marine Science 17:330—337.

, and R. S. MacKenzie. 1971. Distribution of phytoplankton species at the western margin of

the North Atlantic Ocean.—Bulletin of Marine Science 21:603-612.

Johnson, P. W., and J. McN. Sieburth. 1979. Chroococcoid cyanobacteria in the sea: A ubiquitous
and diverse phototrophic biomass.—Limnology and Oceanography 24:928—935.

Malone, T. C. 1971. The relative importance of nannoplankton and netplankton as primary producers
in tropical oceanic and neritic phytoplankton communities.—Limnology and Oceanography
16:633-639.

Marshall, H. G. 1969. Phytoplankton distribution off the North Carolina coast.—American Midland

Naturalist 81:241—257.

. 1971. Composition of phytoplankton off the southeastern coast of the United States.—Bul-

letin of Marine Science 21(4):806-825.

. 1981. Occurrence of bluegreen algae (Cyanophyta) in the phytoplankton off the southeastern

coast of the United States.—Journal of Plankton Research 3:163—-166.

. In Press. Phytoplankton assemblages within the Chesapeake Bay plume and adjacent waters

of the continental shelf.—Estuarine, Coastal and Shelf Science.

, and M. S. Cohn. 1981. Phytoplankton community structure in northeastern coastal waters

of the United States. I. October 1978.—NOAA Technical Memorandum NMFS-F/NEC-8,

57 p.

McCarthy, J. J.. W. R. Taylor, and J. Loftus. 1974. Significance of nanoplankton in the Chesapeake
Bay estuary and problems associated with the measurement of nanoplankton productivity.—
Marine Biology 24:7-16.

Parke, M., and P. S. Dixon. 1976. Checklist of British marine algae. Third Revision.—Journal of the
Marine Biological Association United Kingdom 56:527-594.

Waterbury, J., S. Watson, R. Guillard, and L. Brand. 1979. Widespread occurrence of a unicellular,
marine, planktonic, cyanobacterium.—Nature 277:293-294.

Department of Biological Sciences, Old Dominion University, Norfolk, Virgin-
la 23508.

PROC. BIOL. SOC. WASH.
95(1), 1982, pp. 114-115

NOMENCLATURAL STATUS OF THE FORAMINIFERAL
GENUS CUBANELLA SAIDOVA, 1981

Richard W. Huddleston and Drew Haman

The Carpenteriinae, as defined by Saidova (1981) contains Carpenteria Gray,
1858, Haerella Belford, 1960, Carpenterella Bermudez, 1949, and Cubanella Sai-
dova, 1981. Carpenterella Bermudez is a junior homonym of Carpenterella Col-
lenette in Collenette and Hale-Carpenter 1933, and was renamed Bermudezella
by Thalmann (1951). Loeblich and Tappan (1964) considered Bermudezella and
Haerella junior synonyms of Carpenteria.

Saidova (1981) established the genus Cubanella and selected Carpenteria cu-
bana (Cushman and Bermudez, 1936) as the type-species. Originally this species
was described as the type and only member of Neocarpenteria by Cushman and
Bermudez (1936), and they differentiated Neocarpenteria from Carpenteria based
on its large, semicircular aperture with distinct lip. Loeblich and Tappan (1964)
noted that the aperture of the type-specimen of Neocarpenteria cubana was an
irregular artifact of the broken final chamber. Loeblich and Tappan (1964:C708)
further added, ‘‘As the type species of Carpenteria does not grow upward into
a cylindrical form and the apertural distinction is nonexistent, the name Neocar-
penteria is a junior synonym [of Carpenteria].’’ Cubanella Saidova is a junior
objective synonym of Neocarpenteria (International Code of Zoological Nomen-
clature, ICZN, Article 61b) with Neocarpenteria a junior subjective synonym of
Carpenteria (Loeblich and Tappan, 1964). Cubanella Saidova also is a junior
homonym of Cubanella Pelegrin Franganillo (1926), an arachnid.

Cubanella Saidova 1981, cannot be given a new name (ICZN, Article 60), but
is to be replaced by Carpenteria, the oldest available synonym (ICZN, Article
60a); or if the synonymy of Neocarpenteria and Carpenteria is rejected, by Neo-
carpenteria.

Acknowledgments

We thank K. L. Finger, Chevron Oil Field Research Company for reading this
manuscript and helpful comments, and Chevron Oil Field Research Company for
permission to publish.

Literature Cited

Belford, D. J. 1960. Upper Cretaceous Foraminifera from the Toolonga calcitutite and Gingin Chalk,
western Australia.—Australian Bureau of Mineral Resources, Geology and Geophysics, Bul-
letin 57, 198 pp.

Bermudez, P. J. 1949. Tertiary smaller Foraminifera of the Dominican Republic.—Cushman Labo-
ratory of Foraminiferal Research, Contributions, Special Publication 25, 322 pp.

Collenette, C. L., and G. D. Hale-Carpenter. 1933. New species and description of larva of Lyman-
triidae from Uganda.—Entomology Monthly Magazine 69:258-270.

Cushman, J. A., and P. J. Bermudez. 1936. New genera and species of Foraminifera from the Eocene
of Cuba.—Cushman Laboratory of Foraminiferal Research, Contributions 12(2):27-63.

Gray, J. E. 1858. On Carpenteria and Dujardina, two genera of a new form of Protozoa with attached

VOLUME 95, NUMBER 1 115

multilocular shells filled with sponge, apparently intermediate between Rhizopoda and Porif-
era.—Zoological Society of London, Proceedings 26:266—271.

Loeblich, A. R. Jr., and H. Tappan. 1964. Treatise on Invertebrate Paleontology, Part C, Protista 2,
Chiefly Thecamoebians and Foraminiferida (ed. R. C. Moore).—University of Kansas Press,
Lawrence, 900 pp.

Pelegrin Franganillo, R. P. 1926. Nuevos o poco conocidos de la isla de Cuba.—Boletin Sociedad
Entomologica Espana 9(3-4):42-68.

Saidova, Kh. M. 1981. O sovremennom sosteyanii sistemy Nadvidovykh Taksonov Kajnozojskihh
bentosnykh foraminifer.—Akademiia Nauk SSSR Isstytut Okeanologiia, 73 pp.

Thalmann, H. E. 1951. Mitteilungen uber Foraminiferen.—Eclogae Geologicae Helvetiae, pt.
9(43):221—225.

Chevron Oil Field Research Company, P.O. Box 446, La Habra, California
90631.

PROC. BIOL. SOC. WASH.
95(1), 1982, pp. 116-160

SOME NEW AND OLD SPECIES OF THE PRIMNOID GENUS
CALLOGORGIA GRAY, WITH A REVALIDATION OF
THE RELATED GENUS FANELLIA GRAY
(COELENTERATA: ANTHOZOA)

Frederick M. Bayer

Abstract.—One new and three previously known species of Callogorgia Gray
are described and illustrated by scanning electron microscopy. The diagnostic
characters of the genus are discussed and the species divided between two genera,
for the second of which Gray’s generic name Fanellia is restored to validity.
Keys to the species of both (exclusive of the Atlantic and Mediterranean) are
provided.

Introduction

In the course of reviewing the genera of Gorgonacea, it was found that the
species traditionally assigned to the genus Callogorgia (Family Primnoidae) fall
into two separate groups on the basis of external skeletal ornamentation, loosely
correlated with colonial growth form. Typical Callogorgia, based upon Gorgonia
verticillata Pallas, has pinnate growth form and sclerites varying from externally
smooth to granulated or wrinkled, in a few species with the abaxial scales sculp-
tured by prominent radial crests derived from the marginal ridges of the inner
rim. A second group of species has dichotomous branching (or pinnate branching
so lax that it may appear dichotomous, which 1s here called ‘quasi-dichotomous’)
and sclerites with crowded, radially oriented tubercles extensively coalesced as
ridges. This group of species is now restored to generic status under the name
Fanellia, proposed for Primnoa compressa Verrill by J. E. Gray (1870:46). This
genus as now constituted occurs from the Bering Sea and Gulf of Alaska south
and west to Hawaii and New Caledonia, across the Pacific to Japan and Indonesia.
It is not known from the eastern Pacific, from either side of the Atlantic or from
the Mediterranean, and has not been reported from the Indian Ocean. Species of
Callogorgia occur in the Mediterranean Sea, on both sides of the Atlantic Ocean,
from the west coast of the Americas westward to Indonesia, and throughout most
of the Indian Ocean. The few Callogorgia species reported from the Southern
Ocean (C. antarctica, C. kuekenthali, C. nodosa and possibly C. ventilabrum),
all with dichotomous growth form, are referable to still another genus, Ascolepis
Thomson and Rennet, having sculpture completely different from that of Fanellia.

Methods

The Scanning Electron Micrographs (SEM) of intact polyps were made with
a Cambridge Stereoscan model S4-10, and the sclerites were examined and pho-
tographed with a Coates and Welter model 106 field-emission instrument.

For examination by SEM, sclerites were dissociated and cleaned of surrounding
tissues by commercial 5.25% sodium hypochlorite bleach, followed by weak (3%)

VOLUME 95, NUMBER 1 LA)

hydrogen peroxide and thorough washing in water. After a rinse in 100% ethanol,
the bulk sample was dried and subjected to an oxygen plasma for about 4 hours,
before mounting with PVA emulsion adhesive on 10-mm coverglasses that can
be affixed to standard Cambridge SEM stubs, or held in a custom-made spring-
clip holder compatible with the Cambridge stage. Specimens were precoated with
carbon and sputter-coated with gold/palladium.

Samples for examination as whole mounts were selected for undamaged con-
dition using a conventional stereomicroscope. After removal of superficial tissue
by a very brief immersion in sodium hypochlorite solution followed by hydrogen
peroxide and thorough rinsing in 70% ethanol, air-dried samples of suitable length
(no more than 2 cm) were mounted vertically in the center of standard Cambridge
stubs by means of PVA emulsion adhesive. After appropriate coating with carbon
and gold/palladium, these preparations were placed in the Cambridge series 200
microanalytical stage using an adapter specially made to accommodate the stan-
dard Cambridge stub. The smaller Cambridge series 100 stage will not accept
samples of this size. The stage was tilted to a full 90° for examination of such
wholemounts, and stereo pairs prepared by rotation (8°) rather than by tilting, in
order to preserve the original orientation more closely.

Taxonomic Characters

The principal characters available for the classification and identification of
primnoids are:

1 Colonial form and manner of branching.

2 Size and arrangement of polyps.

3 Number, arrangement, form and ornamentation of sclerites on the polyps and,
to a lesser extent, on the coenenchyme.

Colonial form.—Although ‘‘pinnate’’ and ‘‘dichotomous’’ branching have been
employed as key characters, the two forms intergrade, and where they do so they
are ambiguous. In true pinnate growth form, only branches of the first order are
produced; the first-order branchlets are terminal ‘‘pinnae’’ that rarely subdivide,
except to produce a lateral plume that is structurally identical to the main one.
In the dichotomous growth form, branching proceeds to several orders, each
branch repeatedly forking into two. If the first-order branchlets in a pinnate col-
ony are widely spaced and are not consistently ‘‘terminal’’ but produce second-
and third-order branchlets, the branching pattern simulates dichotomy. This con-
dition is illustrated in Figs. 15 and 18, and is here called ‘‘quasi-dichotomous.’’

Size and arrangement of polyps.—The distribution of polyps in the colony
appears to be a consistent character at generic and specific levels. The verticillate
arrangement is common in the family and, in genera such as Primnoella, Calyp-
trophora, and Narella, is never departed from. In all known species of Callo-
gorgia save one, the polyps are in whorls, and the one exception differs in so
many other regards that it probably does not belong in Callogorgia at all. In
Plumarella, the polyps may be arranged either in tight spirals all around the
branchlets, or biserially (but not in opposite pairs, which really are ‘‘whorls’’ of
two). In Thouarella, various species are described with polyps paired, in whorls,

118 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

and in loose, irregular spirals even within one subgenus of this large and per-
plexing genus. It is probable that when all the species have been reviewed in
modern terms, a generic realignment in which the arrangement and spiculation
of the polyps is more consistent will emerge.

The direction in which the polyps face is quite consistent. Except in the genus
Primnoa, polyps face upward in all genera in which they are not in whorls; the
polyps of Primnoa face downward, apart from occasional strays. In Primnoella,
Callogorgia, and Fanellia they always face upward, again except for strays
(which may be induced by the presence of epizoites, such as echinoderms). In
Calyptrophora they always face upward, in Paracalyptrophora, Arthrogorgia,
and Narella always downward, and in Candidella, Parastenella, Pterostenella,
and the verticillate species of Thouarella, directly outward or slightly upward.

Sclerites.—Although irregular tuberculate rods and spindles occur in the Prim-
noidae and may be the original sclerite form in this family, they are of only
sporadic occurrence and the predominant forms are flat scales or plates, some-
times very thick. On the body of the polyps, these scales are basically arranged
in 8 longitudinal rows as well as in transverse circlets, of which the uppermost
constitutes in most genera an operculum of 8 triangular scales distinctly differ-
entiated from the rest of the body scales below them. The sizes and shapes of
the sclerites are influenced by the extent of inward curvature achieved by the
polyps during contraction. A special terminology of sclerites in various positions
on the polyps has developed over the years, differing even from genus to genus,
depending upon the degree of modification that the skeletal armature has under-
gone. Because these terms are generally used in keys and taxonomic descriptions,
those applicable to forms with strongly in-turned polyps, such as Callogorgia,
are briefly explained.

Because of the inward curvature of the polyps, those rows on the side of the
body turned toward the axis are called adaxial, those on the side facing outward
abaxial, and the intermediate ones outer lateral and inner lateral. Owing to the
bilateral symmetry of the octocoral polyp, the scale rows are paired, so the
members of the pairs lie on opposite sides of the body, with the ‘‘sagittal’’ plane
passing between the members of the abaxial and adaxial scale rows. The scales
of the transverse circlet immediately below and surrounding the operculars are
termed marginal, but no special terminology applies to those below. To save
Space in print, the scale rows are designated by abbreviations (Abax = abaxial;
OL = outer lateral; [IL = inner lateral; Adax = adaxial) and the individual scales
are numbered from the operculum downward, as shown in Fig. 1.

In Callogorgia the outer surface of the sclerites may be smooth, wrinkled, or
ornamented by sharp granules often aligned in radial or reticulating rows (see
Figs. 9, 10). The inner surface of the body scales always has a more or less
smooth distal margin marked by radial ridges that may be low, or high and crest-
like, throwing the edge into dentations (see Figs. 7, 8), and sometimes extending
outward as tall crests on the distal outer surface of the scales (see Figs. 2—5).

In Fanellia the outer surface is covered by crowded, fluted or serrated tubercles
commonly coalescing to form radial ridges (see Figs. 13, 14; 16, 17; 19-22; 25,
26; 28, 29). In both genera, the central inner surface of all scales is covered by
crowded, complex tubercles that serve to anchor the sclerites in the mesogloea.

VOLUME 95, NUMBER 1 119

Fig. 1. Arrangement and terminology of body sclerites, diagrammatic: A, Fanellia with both
outer-lateral and inner-lateral rows reduced in number; B, Callogorgia with outer-lateral rows well
developed, inner laterals reduced. Abax = abaxial; OL = outer lateral; IL = inner lateral; Adax =
adaxial (not visible in side view). O = opercular scale; 1 = marginal; 2— = remaining body scales
numbered proximad.

Callogorgia Gray, 1858

Gorgonia.—Pallas, 1766:160 (part).—Linnaeus, 1767:1289 (part).—Ellis and So-
lander, 1786:67 (part).

Muricea.—Dana, 1846:675 (part).

Prymnoa.—Ehrenberg, 1834:357 (part).

Primnoa.—Milne Edwards and Haime, 1857:139 (part)—von Koch, 1878:457;
1887:85.

Callogorgia Gray, 1857 [1858]:286 (type-species, Gorgonia verticillata Pallas,
1766, by monotypy).—Bayer, 1961 [1962]:296 (part).—Carpine and Grasshoff,
nO75: 102.

Calligorgia Gray, 1870:35 (unjustified emendation).—Studer, 1878:51.

Xiphocella Gray, 1870:56 (type-species, Gorgonia verticillata: sensu Esper,
1797:156, by monotypy).

? Callicella Gray, 1870:37 (type-species, Callicella elegans Gray, 1870, by mono-
typy).

Caligorgia Wright and Studer, 1889:75 (unjustified emendation).—Versluys,
1906:55.—Kukenthal and Gorzawsky, 1908:19.—Kinoshita, 1908:34.—Nutting,

120 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1908:574.—Kukenthal, 1912:320(?); 1915:146; 1919:362; 1924:267.—Deich-
mann, 1936:158.

Description. —Colonies of moderate (30 cm) or large (over 1 m) size, branched
in one plane, pinnate, with branchlets usually alternating, rarely opposite (1
species), new branchlets originating only apically, not between pre-existing
branchlets, sometimes openly pinnate or quasi-dichotomous, rarely dichotomous
(1 species?). Polyps in whorls of 3-6 or more, directed toward apex of branches,
in some species beginning distally as pairs and increasing in numbers proximad,
bent inward toward the axis, the adaxial side more or less closely appressed to
the axial coenenchyme. Sclerites of the polyps in the form of scales curved to
conform with the contours of the body, placed in longitudinal rows of which only
the two abaxials are complete, the lateral and adaxial rows with reduced numbers
of scales, in some species absent adaxially except for the operculars. Opercular
scales triangular, shape abruptly different from those of marginals, which do not
fold inward over the operculars. Tentacles without sclerites in most species, a
few with minute rods. Coenenchymal sclerites in the form of plates, from broadly
polygonal or ovate to strongly elongate. Axis heavily calcified, irregularly grooved
longitudinally, yellowish, brownish or greyish, often with metallic or iridescent
sheen.

Discussion.—This genus, established by J. E. Gray (1858) for the eastern At-
lantic and Mediterranean Gorgonia verticillata Pallas, 1766, is widespread and,
apart from some trans-Atlantic confusion of the type-species (see Carpine
1963:30; Carpine and Grasshoff 1975:102), has been comparatively free of prob-
lems. However, a number of species have been ascribed to it erroneously, and
examination of a wide array of species reveals that they fall into rather well-
defined groups on the basis of spicular ornamentation.

Kukenthal (1924:267) characterized the genus as follows (translation mine):
Colonies abundantly branched and mostly in one plane, sometimes pinnate with
alternate or opposite terminal branchlets, sometimes more dichotomous. The
polyps occur in whorls, only on the larger stems scattered as well. Their adaxial
wall is more or less naked. The scales of the polyps are set in longitudinal rows,
of which the adaxials are always reduced; they are mostly strong and conspicu-
ous, beset on the inner surface with numerous closely placed small warts, on the
outer surface mostly with thornlike projecting ridges or other sculpture. The
opercular scales are well developed abaxially and distinctly pointed, becoming
smaller adaxially. The marginal scales do not overreach the operculars and are
not movable inward. The coenenchymal scales are unlike those of the polyps, not
imbricated but set side by side and mostly quite thick, commonly elongated.

It is clear from published descriptions that Caligorgia antarctica Kukenthal
(1912:321), Caligorgia nodosa Molander (1929:60), and Caligorgia ventilabrum
Studer sensu Gravier (1914:85) actually belong to Ascolepis Thomson and Rennet
(1931:20), which I regard as a valid genus (Bayer 1981:936). The growth form of
all of these species is quite unlike the pinnate colonies of most species of Cal-
logorgia.

Versluys (1906:83), in his exemplary monograph on the Primnoidae of the Si-
boga Expedition, recognized that the species of Callogorgia reviewed by him fall
into groups. Unfortunately, he grouped the species first on the basis of branching,

VOLUME 95, NUMBER 1 121

which, as can be seen in other genera of Primnoidae, is an unreliable character,
partly because it is difficult to distinguish pinnate branching, if widely spaced,
from dichotomous, and partly because both ‘‘pinnate’’ and ‘‘dichotomous”’
branching can occur in a single genus.

However, among the dichotomously branched species, Versluys (1906:84) clear-
ly recognized the distinctively different kind of sculpturing characteristic of the
sclerites of C. compressa and C. tuberculata. The subsequent description of two
other species (C. granulosa, C. aspera) from Japan by Kinoshita (1907, 1908a) and
one from the Marshall Islands by me (Bayer 1949:207), together with the discov-
ery among the Albatross Philippine octocorals of yet another species, all with the
same kind of external ornamentation, makes it clear that this character, not the
colonial growth form, provides a reliable basis for grouping of species. Further,
neither in the literature nor among the specimens examined have I found any
examples of sculpture intermediate between the tuberculata-compressa type and
the verticillata-flabellum type. Consequently, these two groups are discontinuous
and merit full generic rank. A new generic name is not required for the tuber-
culata-compressa complex as Gray (1870:46) already has proposed for C. com-
pressa Verrill a nominal genus Fanellia, which is here restored to good standing.

Although the remaining species of Callogorgia have generally similar sculpture,
two sub-types can be recognized among them: (1) those with the scales more or
less smooth externally, sculptured at most by smooth granules, usually scattered
but sometimes merging to form wrinkles that may anastomose into reticular or-
namentation (sertosa Wright and Studer, verticillata Pallas, grimaldii Studer,
pennacea Versluys, joubini Versluys, similis Versluys, minuta Versluys, affinis
Versluys, kinoshitae Kukenthal, gilberti Nutting, ventilabrum Studer, gracilis
Milne Edwards and Haime); and (2) those with the body scales sculptured ex-
ternally by prominent radial crests that are outward extensions of the radial ridges
developed around the inner distal margin of the body scales in most species, if
not all (fabellum Ehrenberg, weltneri Versluys, robusta Versluys, cristata Au-
rivillius, verticillata sensu Deichmann). Even though these two kinds of sculpture
are usually quite distinct, there is some overlap. In some (but not all) specimens
of C. gracilis (Milne Edwards and Haime), the inner marginal ridges of the abaxial
marginal scales are reflected outward onto the outer surface as low ridges con-
verging toward the nucleus of the scale. These ridges have a quite different aspect
from those of C. flabellum and related forms (robusta, weltneri, cristata), but
they seem to be identical structurally. Therefore, there seems to be no infallible
basis for subdividing those species into two genera, or even subgenera.

The disposition of Ascolepis Thomson and Rennet, 1931, is equivocal. Both of
the original species, A. splendens and A. spinosa, have sclerites of unusual
form—not so distinctly cuplike as originally described and illustrated from ob-
servation with the light microscope, but certainly with a concave outer part more
or less clearly set off from a tuberculated base. The same is true of Caligorgia
nodosa Molander and the specimens reported as C. ventilabrum by Gravier,
appears to be true also of C. antarctica Kukenthal, and could be true of C.
ktikenthali Molander. If C. antarctica represents an intermediate between Cal-
logorgia and Ascolepis, thus making the latter a junior synonym, then the genus
Callogorgia will include species that do not remotely resemble the majority of
species of Callogorgia. Because of the difficulty in defining Callogorgia that

122 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

would result from merging Ascolepis with it, the preferable course is to maintain
both as distinct.

Other problems with the definition of Callogorgia remain to be solved, notably
the case of those Antarctic species of C. antarctica growth form in which the
regular alignment of abaxial and lateral body scales is completely disrupted.
Whether these merit separate recognition, or must be accommodated in a more
broadly defined genus Callogorgia is a question that must await further study.
For the time being, I propose to recognize Callogorgia, Fanellia, and Ascolepis
as separate genera.

Key to Indo-Pacific Species of Callogorgia!

1(20). Branching typically pinnate.
2(3). Branches strictly opposite .............. C. formosa Kukenthal, 1907
3(2). Branches alternate.
4(7). Outer-lateral scale-rows well developed.
5(6). Polyps 1.3—1.5 mm tall, whorls separated by conspicuous intervals,
SW ONIN, ye ca tas oe coheed ae Cae C. sertosa Wright and Studer, 1889
6(5). Polyps larger, up to 2 mm tall, whorls closely placed, 5in 1 cm ...
PR, Pee Wiig a, Oia aCe SERN Maa BE 29 0 0S a al C. kinoshitae Kukenthal, 1913
7(4). Scales in outer lateral rows sharply reduced in number.
8(19). Operculum tall and prominent.
9(16). Apex of opercular scales with a single point.
10(13). 10 scales in each abaxial row.
11(12). Apex of opercular scales prolonged into a rodlike point ...........
LN Ea iN, YR Mrs aie PENT RA C. ramosa Kukenthal & Gorzawsky, 1908
12(11). Apex of opercular scales not prolonged into a rodlike point .......
YG PN ee eae at i IEnen NPIS SIS eee oat ON 8 o C. flabellum (Ehrenberg, 1834)
13(10). 6-7 scales in each abaxial row.
LAGS) Polyps. .2. min tally se chocysnceescgaic eno Seneca ieee C. robusta Versluys, 1906
TSCA). Jolypos wma, ball. 00a satack uve my sled uae C. joubinii Versluys, 1906
16(9). Abaxial opercular scales with 2-4 apical points.
17(18). 5-6 whorls of polyps in | cm; 8-10 scaies in abaxial rows; no inner

lateralisCales: occ eae. eee a ee C. cristata Aurivillius, 1931

18(17). 4-5 whorls of polyps in | cm; 7 scales in abaxial rows; | inner-lateral
SCANS Fle iear ie ols! ceva syth 5h ae pec ae WC ee a C. weltneri Versluys, 1906
19(8) Operculumi lows ans0 wcuus ey aoe eo C. pennacea Versluys, 1906

20(1). Branching dichotomous or quasi-dichotomous.
21(26). Outer-lateral scale rows well developed.

PA(O3) 8) scalesuinueach abaxda lajOwa sere eee C. ventilabrum Studer, 1878
23(24). 9 scales in each abaxial row C. laevis Thomson and Mackinnon, 1911?
2A@S). lO;scales imzeach abaxial rows - 0.8 545. C. versluysi Thomson, 1905
25(22). 12-13 scales in each abaxial row ............. C. elegans (Gray, 1870)
26(21). Scales in outer lateral rows reduced in number.

27(28). 3 scales in each outer-lateral row ........... C. indica Versluys, 1906

1 Modified from Kukenthal (1919).
2 Examination of the type-specimens in the Australian Museum since this manuscript went to
press has enabled me to determine that this species does not belong to Callogorgia but to Primnoella.

VOLUME 95, NUMBER I 123

28(27). 1 or 2 scales in each outer-lateral row.

29(32). 5 scales in each abaxial row.

30(31). 2-3 polyps per whorl, rarely 4; 8-9 whorls in 1 cm; body scales with
lopranmmancimal TGSCS 2.060042. eae ee ee C. minuta Versluys, 1906

31(30). 3-4 polyps per whorl, sometimes 5; 9-11 whorls in 1 cm; body scales
with distal margins strongly reflexed, exposing high crestlike ridges
I ec ON: ks os loa Ha bate Pew Be C. chariessa, nN. sp.

32(29). 7 scales in each abaxial row.

33(34). Body scales with strong marginal ridges; 4-5 polyps in each whorl,

SE NOmmmoms te leCM’ so... 2. ee abe ee ee C. affinis Versluys, 1906
34(33). Body scales with weak marginal ridges; 2, usually 3 polyps in each
Winggieme-oo WHOTlS i 1 CM 2... ee ce ee C. similis Versluys, 1906

Callogorgia flabellum (Ehrenberg)
Figs, 2,3

Prymnoa flabellum Ehrenberg, 1834:358 (locality not given).

Caligorgia flabellum.—Versluys, 1906:69, figs. 75-78, pl. 5, fig. 13, pl. 6, fig. 14
(Kei Islands, 5°28.4’S, 132°0.2'’E, 204 m).

not Caligorgia flabellum.—Nutting, 1912:60 (=C. aspera Kinoshita, 1908a = C.
tuberculata Versluys, 1906).

Material examined.—Moluccas, off Makian, west of Halmahera, 0°15’00’N,
127°24'35"E, 545 m, sand and coral, Albatross sta. DS621, 28 November 1909:
several large branches without base, and smaller detached branches, USNM
57545 (Figs. 2, 3).

Description.—See Versluys, 1906:69.

Discussion.—As Versluys (1906:69) has given a full description of the Siboga
material, description of the present material would be superfluous. However, the
validity of Versluys’s identification of the Siboga material rests solely on his own
comparison with a badly preserved fragment of Ehrenberg’s type in the Berlin
Museum, as Wright and Studer did not compare the Challenger specimen with
Ehrenberg’s type as Versluys thought (1906:71), but with Gray’s type of Callicella
elegans (Wright and Studer, 1889:79).

The specimen taken by the Albatross off Makian, west of Halmahera in the
Moluccas, agrees satisfactorily with Versluys’s account of the Siboga specimens
from the Kei Islands. The characters that are diagnostically important are: 8-10
abaxial scales in each row, with strong radial ridges extending from the inner
marginal ridges; 3 outer laterals, 2 inner laterals, | adaxial; operculars tall, tapered
to a single blunt point; abaxial scales sculptured with strong radial crests that
extend distally as marginal serrations, most prominent on the distal sclerites,
decreasing in strength proximad until on the basalmost one or two they are re-
duced to low ridges or merely conspicuous wrinkles. The cortical plates are
irregularly elongate, commonly with a central outer projection and crowded,
pointed tubercles that are more or less extensively joined to form irregularly
meandering low ridges or wrinkles.

The present material differs from Versluys’s description to a small degree,
which records 2 outer laterals and | inner lateral but, in fact, his drawing (Fig.
85) suggests 3 outer laterals and 2 inner laterals, as is usually the case in the

124 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Callogorgia flabellum (Ehrenberg), USNM 57545: left and right lateral views of polyp.
SEM stereo pairs, x40.

specimen before me. As can be seen from the accompanying SEM photos (Fig.
2), the sculpture of the body scales obscures the distinction between individual
scales and renders counting and drawing by camera lucida uncertain. The shapes
and positions of scales shown in Versluys’s drawing make it difficult to interpret
the scales labeled ‘G’ as outer laterals. Their location requires them to be inner
laterals, hence 2 in number, so the adaxial part of the 3 distalmost abaxial scales
can only be 3 outer laterals.

Callogorgia flabellum is very similar to C. cristata in aspect, difficul to dis-
tinguish with the dissecting microscope. The most reliable recognition features
are: (1) the presence of 3 OL, 2 IL, and 1 Adax; (2) the single points on the
abaxial operculars; (3) the extension of the inner marginal ridges of the abaxial

VOLUME 95, NUMBER 1 125

Fig. 3. Callogorgia flabellum (Ehrenberg), USNM 57545: opercular, adaxial marginal, abaxial,
outer-lateral sclerites of polyp. Scale = 0.1 mm.

126 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Nin peng
(Vb

Fig. 4. Callogorgia cristata Aurivillius. Top, USNM 57544; bottom, USNM 44179, part of type
in Zoological Museum, University of Uppsala: intact whorls of polyps. SEM stereo pairs, X27.

scales onto the outer surface as strong radial crests; and (4) the persistence of
external sculpture, though reduced, on the proximal abaxial scales.

Callogorgia cristata Aurivillius
Figs. 4, 5

?Caligorgia weltneri Versluys, 1906:73, figs. 80-82, pl. 4, fig. 10 (southwest of
Waigeu, between Jef Fam and Gagi, 0°29.2'S, 130°05.3’E, 469 m).—Kukenthal,
1924:274.

Caligorgia cristata Aurivillius, 1931:262, fig. 52, pl. 6, fig. 1 (Japan, Goto Is.,
west of Kyushu, 160 m).

VOLUME 95, NUMBER 1

Fig. 5. Callogorgia cristata Aurivillius, USNM 57544: opercular, abaxial, outer-lateral, and mod-
ified opercular scales of polyp. Scale = 0.1 mm.

128 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Material examined.—Japan, Goto Islands, off Pallas Rock, 32°15’N, 128°12’E,
160 m, coral bank, 15 May 1914, coll. S. Bock: small branch of type, USNM
44179 (Fig. 4, bottom). |

Moluccas: off Ternate, 0°49'30"’N, 127°25’'30’E, 240 m, coral; Albatross sta.
D5617, 27 November 1909: large branch and detached smaller branches, USNM
57544 (Fig. 4, top).—off Makian, 0°15’00°N, 127°24'35”E, 545 m, sand and coral;
Albatross sta. DS5621, 28 November 1909: one pinnate branch without base,
USNM 60285.

Description.—See Aurivillius, 1931:262. Compare Versluys, 1906:73.

Discussion.—The specimens collected by the Albatross from Ternate and Mak-
ian, both off the west coast of Halmahera in the Moluccas, agree well with Au-
rivillius’s account of C. cristata, and with Versluys’s description of C. weltneri.
The principal difference between C. cristata and C. weltneri cited by Aurivillius
(1931:266) is the *“‘somewhat greater number of polypal spicules, amounting to
8—9(-10) as against only 7’ in weltneri. The present specimens tend to have
somewhat more scales in the abaxial rows than was usual in C. cristata, but this
character is subject to variation in all species of Callogorgia. Direct comparison
by SEM of USNM 57544 (Fig. 4a) with type-material of C. cristata (Fig. 4b)
reveals no other significant difference, so they can safely be considered conspe-
cific. Even though Versluys (1906:74) reports no variation from 7 abaxials in C.
weltneri, deviations almost certainly occur. A difference of 1 in the prevalent
number of abaxial sclerites is a weak justification for specific separation so it is
very likely that cristata and weltneri represent a single species. However, as
type-material of C. weltneri has not been examined in this context, C. cristata
is maintained. The inner basal angle of the inner-lateral opercular scales (IL-0)
is in many cases expanded as a rounded lobe that extends into the position that
would be occupied by the inner-lateral marginal scale (IL-1) if such a sclerite
were developed (Fig. 5). This condition was not noticed by Aurivillius, but its
inconsistent development may have led to its oversight.

Callogorgia chariessa, new species
Figs. 6-8

Material.—One pinnately divided branch; Tawitawi Group, Sulu Archipelago,
5°18'10"N, 120°02’5S"E (Tocanhi Point S.27°E, 2.1 miles), 49 fath. (90 m),
coral sand and shell; Albatross sta. D5153, 19 February 1908. Holotype, USNM
58398.

Description.—Callogorgia with alternate pinnate branching in one plane, the
nodes 5-10 mm apart and the undivided terminal branchlets about 2 cm long,
producing the open type of pinnate branching that easily can be mistaken for
dichotomous. Polyps in whorls of 3—S, commonly 4; in the terminal branchlets,
9 whorls commonly occur in 1 cm, uncommonly as many as 10 or 11. Diameter
of undivided terminal branchlets 0.5 mm exclusive of polyps; axis alone about
0.3 mm in diameter, tapering to a hair-fine apex. Axis heavily calcified, longitu-
dinally striated, creamy yellow with golden iridescence.

Polyps with only the two abaxial rows of scales well developed, 4 or 5 scales
in each; one large outer lateral on each side, although this is occasionally absent,
its place being occupied by the distalmost abaxial which then is sufficiently broad-

VOLUME 95, NUMBER 1 129

Fig. 6. Callogorgia chariessa, new species; holotype colony, USNM 58398. Scale in cm.

ened to cover the space; distal margins of the body scales strongly reflexed out-
ward exposing the marginal radial ridges, which are high and crestlike. Opercular
scales roughly triangular, with serrated edges, rounded basal angles and 1-4 ser-
rated ridges on both outer and inner surfaces of the blunt apex; adaxial and inner
lateral operculars conspicuously smaller than the abaxials and outer laterals. Ex-
posed surface of scales with sparse granular sculpture, partly aligned in irregular
radial rows, inner surface with compound tubercles. Cortical scales large, polyg-
onal, closely interlocking by their marginal serrations.

Comparisons.—This species is similar to C. minuta and C. similis (Versluys,
1906:76, 78); the latter, in turn, is ‘‘im Habitus der C. ventilabrum ausserst ahn-
lich’’ (Versluys 1906:74, 76). C. ventilabrum, which originally was collected in
162 m north of New Zealand by the Gazelle Expedition, has 7 scales in the abaxial
rows (as determined by Versluys 1906:75, who examined a part of the type col-
ony) compared with 5 or fewer in C. chariessa. The inner distal edges of the
body scales of C. ventilabrum have sharp radial ridges but they are comparatively
low and not conspicuously exposed by the outward reflexion of scale margins as
in C. chariessa.

Callogorgia chariessa resembles C. minuta even more closely, but the whorls
tend to be more closely placed (as many as 10 or 11 per cm, though commonly
9) than in that species (8 or 9 per cm), and to consist of a larger number of polyps
(commonly 4, sometimes 3 or 5, compared with 2 or occasionally 3 but rarely 4
in minuta). The margins of the body scales are strongly reflexed in C. chariessa,
conspicuously exposing the marginal ridges to view. In this regard, C. affinis
approaches C. chariessa more closely, but the polyps have 7 scales in the abaxial
rows as opposed to 5 (or even 4) in C. chariessa, and the margins of the body
scales are not so conspicuously reflexed.

130 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 7. Callogorgia chariessa, new species; holotype, USNM 58398: abaxial and right lateral
views of polyp. SEM stereo pairs, 67.

When more material becomes available for an assessment of variation, it may
prove that all of these basically similar forms belong to a single variable species,
but for the present time it is preferable to call attention to the differences between

them.
Callogorgia formosa Kukenthal
Figs. 9-10

Caligorgia formosa Kukenthal, 1907:208 (Sudwestlich von Gross-Nikobar, 362
m); 1919:366, figs. 155-159, pl. 30, fig. 1, pl. 40, fig. 47; 1924:269, fig. 153.
Primnoella indica Kukenthal, 1907:210.

Material examined.—Hawaiian Islands: north Bank of Necker Island, 250 fath.

VOLUME 95, NUMBER 1 131

Fig. 8. Callogorgia chariessa, new species; holotype, USNM 58398: opercular, adaxial marginal,
abaxial and outer-lateral sclerites of polyp. Scale = 0.05 mm.

(458 m), Townsend Cromwell 76-06-73, sta. 4, 15 October 1976; large colony
broken into 4 pieces: USNM 60692.—Off Necker Island, 23°39'N, 164°31'W,
80-162 fath. (146-296 m), Townsend Cromwell sta. 6, 16 October 1976; two badly
damaged pinnate fragments: USNM 60291.

Description.—A large, plumose colony, broken in 4 pieces measuring 29.5,
18.5, 12.2 and 8.3 cm, corresponds in all essentials with the description and
figures of C. formosa given by Ktkenthal (1919). Evidently broken off well above
the holdfast, the intact colony must have approached | m in height. It is a rather
stiff and brittle pinnate plume with strictly opposite, undivided branches up to
14.5 cm long, the distal ones rather quickly decreasing in length toward the apex.

132 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 9. Callogorgia formosa Kukenthal, USNM 60692: left lateral and abaxial views of polyp.
SEM stereo pairs, x38.

The polyps occur on the branches in rather well-separated whorls of 3 or 4 dis-
tally, increasing proximad to 5, usually 4 in 1 cm; they are in whorls also on the
main axis near its apex, but this arrangement is soon lost and only widely scat-
tered individuals persist on the lower parts of the trunk. The size, shape, and

=

Fig. 10. Callogorgia formosa Ktikenthal, USNM 60692: sclerites of coenenchyme, three outer,
one inner view; opercular, adaxial marginal, abaxial and outer-lateral sclerites of polyp. Scale =
0.1 mm.

VOLUME 95, NUMBER 1

134 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

armature of the polyps is essentially as described by Ktikenthal (1919). He makes
the point (1919:367) that only the distalmost abaxial scales are wider than high,
the rest being rounded or trapezoidal in outline. However, judging by his drawing
(op. cit.: fig. 155), it appears that the proximal abaxial scales may have suffered
some breakage, as is the case in the present specimens. When intact, the proximal
abaxial scales are broad and tapering, extending well around the sides of the .
polyp. The outer-lateral rows are unusually well developed for the genus, com-
posed of 6 or more scales. Except for the distal scales, it is difficult to confirm
Kukenthal’s counts for the inner-lateral and adaxial rows, as the proximal scales
imperceptibly merge with the coenenchymal sclerites, which in full development
are long, irregular spindles sometimes with blunt lobes. The body scales have
only fine granulation externally, but no trace of radial ridges or crests; internally,
they are covered with crowded complex tubercles of usual form, and the distal
margins are sharply and conspicuously toothed. The tentacles are devoid of scler-
ites.

Relationships.—Kukenthal (1919:369) considered this species so unusual that
it might merit a new genus distinct from Callogorgia. Although he considered
that in form of the polyps C. formosa conformed with other species of Callo-
gorgia, he had actually mistaken detached branches for a new species of Prim-
noella (Kukenthal, 1907:210, 1919:368). This demonstrates the difficulty of even
generic identification because of the ambiguity of many taxonomic characters in
the Octocorallia.

Distribution.—In view of its type-locality in the Indian Ocean off Great Nicobar
Island, the discovery of C. formosa in Hawaiian waters is of no little interest. Its
distribution therefore parallels that of Fanellia tuberculata, first found off the
Sulu Islands and now reported from Hawaii, but it has not been reported even
as a synonym from intermediate localities as has F. tuberculata. It is probable
that this large distributional gap results only from inadequate sampling. The other
species of Callogorgia known from Hawaii, C. gilberti Nutting, is more closely
related to C. kinoshitae Kiikenthal from California and C. verticillata (Pallas) sensu
Deichmann from the Caribbean than to any species from the Indo-west Pacific.

Fanellia J. E. Gray, 1870

Fanellia J. E. Gray, 1870:46 (type-species, Primnoa compressa Verrill, 1865; by
monotypy).
Caligorgia.—Auct. (in part).

Diagnosis.—Primnoid colonies branched in dichotomous or lax pinnate manner
simulating dichotomous, rarely close pinnate, usually but not always in one plane.
Polyps upturned, verticillate, in whorls of 2-12; sclerites of polyps arranged in
longitudinal rows in which the distalmost circle of 8 triangular or lanceolate scales
(operculars) is always present; the two abaxial rows of body scales always fully
developed but the outer-lateral, inner-lateral and adaxial rows are more or less
reduced in number of scales owing to inward curvature of the polyps in contrac-
tion (i.e., the adaxial body wall is more or less naked); tentacles devoid of scler-
ites. Coenenchymal sclerites in the form of thick, closely fitting, rounded or

VOLUME 95, NUMBER 1 135

polygonal plates presenting a cobblestone-like aspect. Sclerites sculptured exter-
nally by closely set, angular tubercles that may fuse to form serrated ridges
irregularly meandering or more or less distinctly radiating outward from the cen-
ter, internally by very complex, crowded tubercles not regularly aligned.

Comparisons.—This genus closely resembles Callogorgia but is distinguished
by the external sculpturing of the sclerites, which does not intergrade with the
type of ornamentation developed in Callogorgia. The growth form is predomi-
nantly dichotomous, or so openly pinnate that it appears to be dichotomous (here
called ‘quasi-dichotomous’), but only rarely is it closely pinnate (F. granulosa).
In most Callogorgia species it is regularly pinnate, in some becoming more lax,
but in a few species described (e.g., C. antarctica) it is dichotomous as in Fanellia
compressa. However, several of these dichotomous species seem to be referable
to the genus Ascolepis Thomson and Rennet on the basis of sclerite form. If not,
the branching of Fanellia and Callogorgia will be equivalent, in that both pinnate
and dichotomous conditions occur, together with the quasi-dichotomous inter-
mediate. In Fanellia, branching is predominantly dichotomous, in Callogorgia
predominantly pinnate. In Ascolepis, branching varies even more, ranging from
unbranched (nodosa) to typical dichotomous (splendens, antarctica), lax pinnate
or quasi-dichotomous (spinosa) and bottle-brush form (abies).

Geographical distribution.—Fanellia appears to be confined to the Pacific
Ocean, occurring from the Gulf of Alaska and Aleutian Islands to Japan, Indo-
nesia, Marshall Islands, Hawaii, and as far south as New Caledonia. No species
recognizable as belonging to Fanellia has been reported from the Indian Ocean
or from the Southern Ocean. In contrast, Callogorgia occurs in all seas (unless
all of those from the Southern Ocean prove to be species of Ascolepis), and
Ascolepis occurs only in Antarctic and Subantarctic waters.

Key to Species of Fanellia

1(4). Terminal branches long (15-30 cm), branching dichotomous or quasi-
dichotomous; polyps in whorls of 8-12.
2(3). Polyps with 7-8 scales in each abaxial row, 2 in outer-lateral rows ...
PIE iN. sal 8 kk ie gal gS reat Sle, Qeanala. odie F.. fraseri (Hickson)
3(2). Polyps with 10-11 scales in each abaxial row, 5-7 in outer-lateral rows
ne sig 5 baie wis Sree owls wb pele bee a bs F. compressa (Verrill)
4(1). Terminal branches shorter (7 cm or less), branching clearly pinnate or
quasi-dichotomous; polyps in whorls of 2-5.
so). branching distinctly pinnate .................. F.. granulosa (Kinoshita)
6(5). Branching quasi-dichotomous.
7(8). Colonies flabellate, compressed or in one plane; polyps with 5-8 scales
in abaxial rows, | or 2 outer laterals, 1 inner lateral ................
5 ew optebabag ohn een a Ae tO ee F. tuberculata (Versluys)
8(7). Colonies corymbose, not in one plane; polyps with 5 (occasionally 6)
scales in abaxial rows, one wide marginal scale occupying the combined
inner and outer lateral positions on both sides; no separate inner
[08 )) iets 2 SER RAG EOE aeRO Aer A OR Ae a ea re F. corymbosa, N. sp.

136 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 11. Fanellia compressa (Verrill), USNM 60281: complete colony. Divisions of scale are cm.

Fanellia compressa (Verrill, 1865)
Figs. 11-14

Prymnoa verticillaris.—Ehrenberg, 1834:357.
non Gorgonia verticillata Pallas, 1766:177.

VOLUME 95, NUMBER 1 137

: )
Du” } ; £ ' _ f £

Fig. 12. Fanellia compressa (Verrill), USNM 60281: detail of colony. Divisions of scale are mm.

non Gorgonia verticillaris Linnaeus, 1767:1289.—Ellis and Solander, 1786:83.—
Statius Muller, 1775:753.—Houttuyn, 1772:309, pl. 132, fig. 1.

non Callogorgia verticillata.—Gray, 1858:286.

Primnoa compressa Verrill, 1865:189 (Aleutian Islands).

138 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

aN.

’ ay

tA,

EPSP on cin
G

*
E ate
te

view of polyp, x44. SEM stereo pairs.

Fanellia compressa.—Gray, 1870:46 (new genus for Primnoa compressa Verrill).
Caligorgia compressa.—Versluys, 1906:81.—Kukenthal, 1924:276.

Material examined.—Amchitka, Aleutian Islands, 51°32’N, 179°15'W, 278-289
m, K. K. Chew, coll. 2 September 1968: 4 colonies, complete or nearly so, USNM
60281 (Figs. 11, 12, 14).

Bering Sea, 52°05’N, 177°40’E, 100 m, Albatross sta. 3599, 9 June 1894: 1
colony, USNM 57542.

Bering Sea, 52°14’N, 174°13’E, 882 m, Albatross sta. 4781, 7 June 1906: 1
colony, somewhat broken, USNM 57543 (Fig. 13).

Description.—Colonies flabellate, branched in a very loose pinnate manner that
simulates dichotomous (i.e., quasi-dichotomous), lateral branches arising alter-

139

VOLUME 95, NUMBER 1

> SON
m2

outer-lateral sclerites of

>]

USNM 60281: opercular, abaxial

Fig. 14. Fanellia compressa (Verrill),

polyp. Scale

= 0.1 mm.

140 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

nately, mostly 30-40 mm apart, the undivided terminal branches up to 30 cm
long, nearly straight. The polyps are situated around the branches in closely
placed whorls of 8—12 distally, increasing in numbers proximad until in the major
branches there may be 14-16; on the larger branches some of the whorls may be
oblique, some incomplete, resulting in a spiral appearance; 4—6 whorls occur in
1 cm of length. The polyps are about 1.5 mm tall in the contracted state; the two
abaxial rows of sclerites consist of 10-11 scales each, the outer laterals 5S—7, the
inner laterals 2-3, and the adaxials 1. Because of the greater number of inner and
outer lateral scales in F. compressa as compared with F. fraseri, the winglike
lateral expansion of the abaxial scales has room for development only on those
in the more proximal positions, so it occurs only from Abax-7 to 10; the outer
laterals, beginning between OL-4 and 6, depending upon the number of inner
laterals present, also develop lateral expansions, as may one or more of the inner
laterals. All of the body scales are sculptured externally by closely placed, mi-
nutely aculeate ridges irregularly radiating outward from the nucleus. Where ex-
posed, the coenenchymal sclerites are large, closely fitting polygonal plates sim-
ilarly sculptured, but where covered by the contracted polyps they are smaller
tuberculate spheroids. Small, warty spheroids also occur sparsely in the meso-
gloea between the longitudinal stem canals and immediately surrounding the axis
(i.e., the axial sheath).

Comparisons.—This species resembles F. fraseri but seems to grow larger.
The polyps are larger, the lateral scale rows better developed, and the sculpturing
more complex.

Remarks.—Although Verrill’s original specimens lacked coenenchyme, the lo-
cality from which they came makes it probable that they were the species here
described. Prymnoa verticillaris Ehrenberg could have been either this species
or F. fraseri, as Studer (1878:647) pointed out that Ehrenberg’s material was
obtained in the North Pacific by the Rurik Expedition. Ehrenberg’s statement
that the polyps had 8 scales in the abaxial rows (‘‘scutellorum serie dorsuali
longitudinali duplici, trasversis 8’’) makes it slightly more likely that he had fraseri
in hand, although the difficulty of counting scales accurately leaves room for
doubt. The matter is of no nomenclatural significance because Ehrenberg used
the variant spelling employed by Linnaeus for Gorgonia verticillata Pallas.

Fanellia fraseri (Hickson, 1915)
Figs. 15-17

Caligorgia fraseri Hickson, 1915:553, fig. 4, pl. 1, fig. 2 (Albatross and Portlock
banks, Gulf of Alaska, 50-100 fath. on halibut lines); 1917:23.—Kukenthal,
1924:279.

Material examined.—Gulf of Alaska: Albatross and Portlock banks, 50—100
fathoms (92-183 m); two loosely pinnate branches, paratypes, British Museum
(Nat. Hist.) 1962.7.20.821 (figured by Hickson) and 1962.7.20.822 (Fig. 15B).—
Albatross Bank, 100-125 fath. (183-229 m); M/S Dorothy, Alaska King Crab
Investigations, sta. C-45, 28 March 1941; three branches, probably from a single
colony, USNM 51284 (Fig. 15A).

Description.—Like Hickson’s original specimens, the present topotypic ma-
terial consists of branches possibly from a single colony. Two of the branches

VOLUME 95, NUMBER 1 141

9

t

8

6

LUUTACTLLELNAVATLNVATADUSTHVGTAEATHATAU THA

01

Fig. 15. Fanellia fraseri (Hickson): A, USNM 51284; B, paratypes, BM(NH) 1962.7.20.821 (fig-
ured by Hickson) and 1962.7.20.822. Scale in cm.

(one illustrated in Fig. 15A, right) are very loosely pinnate just as Hickson
(1915:553) described the types, but the third (Fig. 15A, left) is more distinctly
pinnate and probably came from a position lower in the colony. The longest
undivided terminal branchlet is 15 cm long. Distally the polyps are set in whorls
of 5-8, increasing to 12 in the lowest parts of the colony preserved; 6—7 whorls
occur in | cm of length. The polyps are about | mm tall in the contracted state;
the two abaxial rows of sclerites consist of 7-8 scales each (although the distinc-
tion between the lowest abaxial scale and the adjoining coenenchymal sclerites
is often so unclear that an accurate count is difficult to make), and the 3rd, 4th

142 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 16. Fanellia fraseri (Hickson), USNM 51284: top, whorl of polyps, x25; bottom, oblique
view of polyp, x42. SEM stereo pairs.

and 5th on each side have inconspicuous to moderate ““winglike expansions’’ (as
observed by Versluys, 1906:81, Fig. 95, for C. tuberculata); the outer-lateral rows
usually have 2 scales each, the inner laterals 1, and the adaxials 1 small scale
(sometimes missing) below each opercular; the larger coenenchymal sclerites are
closely fitting polygonal plates externally covered by irregular, close-set ridges;
where covered by the contracted polyps, they are smaller, tuberculated grains of
irregular shape.

Comparisons.—This species is generally similar to F. compressa, with the
following differences: branching tends to be somewhat closer and therefore more
distinctly pinnate; polyps smaller, fewer scales in the abaxial and outer-lateral

VOLUME 95, NUMBER 1

Fig. 17. Fanellia fraseri (Hickson), USNM 51284: opercular, adaxial marginal, abaxial, outer-
lateral and inner-lateral sclerites of polyp. Scale = 0.1 mm.

144 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

rows; winglike lateral expansions of the body scales weakly or moderately de-
veloped on Abax-3, 4 and 5, and not at all on outer laterals or inner laterals.

Remarks.—The type-lot was obtained by Prof. Arthur Willey from halibut lines
on Albatross and Portlock banks in the Gulf of Alaska, where it was reported as
common. The color in life was described as pink.

Fanellia granulosa (Kinoshita, 1907)

Caligorgia granulosa Kinoshita, 1907:231 (*Sagami- und Kagoshima-See’’);
1908a:37, pl. 2, figs. 13-14, pl. 6, fig. 46 (°Westkuste von Satsuma’’); 1908b:
jell lis ate, Be LOU.

Caligorgia tuberculata.—Kukenthal, 1924:278 (part).

Material examined.—None.

Discussion.—Kinoshita (1908a:40) stated that the sculpture of the sclerites of C.
aspera and C. granulosa is the same, but that the species differ in branching and
arrangement of polyps. Although his very clear photographs (PI. 2, figs. 13, 14)
of C. granulosa show a distinctly pinnate colony, they also show that the distal
parts of branches, if isolated, could easily be interpreted as dichotomous. More-
over, the differences between the polyps of the two are negligible; even the
presence of 1-3 adaxial body scales below the operculum in aspera compared
with ‘‘vereinzelten Schuppen”’ in granulosa is of no significance. In view of the
intraspecific variation in colonial form that can occur under different ecological
conditions, it seems likely that C. granulosa, C. aspera and C. tuberculata are
indeed but a single species as interpreted by Kukenthal. Although there is little
doubt that Caligorgia aspera Kinoshita is a junior synonym of Fanellia tuber-
culata (Versluys), I prefer for the moment to retain Kinoshita’s C. granulosa as
a distinct species, now referred to the genus Fanellia, because no authentic ma-
terial has been available for examination.

Diagnosis.—Colonies pinnately branched.

Fanellia tuberculata (Versluys)
Figs. 18-26

Caligorgia tuberculata Versluys, 1906:80, figs. 95, 96, pl. 6, fig. 15 (Sulu- Inslen,
5°43.5'N, 119°40’E, 522 m).—Kukenthal, 1924:278 (part).

Caligorgia aspera Kinoshita, 1908a:39, pl. 2, figs. 15, 16; pl. 6, fig. 47 (Westkuste
von Satsuma).—Nutting, 1912:61.

Caligorgia flabellum.—Nutting, 1912:60.

Caligorgia pseudoflabellum Bayer, 1949:207, fig. 2a—c, pl. 4, fig. 2 (Bikini Atoll,
off Enyu Pass, 11°29’28”N, 165°31'40"E, 116-120 fath.).

Material examined.—Hawaiian Islands: Middle Bank, 22°47.5'N, 161°02.2’W,
to 22°47.9'N, 161°02.3'W, 382 m, Townsend Cromwell TC78, 11 January 1978;
broken branches, possibly from one large colony, USNM 56791 (Fig. 19).—Kaena
Point, Oahu, 21°35.85'N, 158°24.55'W, 275-445 m, “‘Sango XII’ haul 1, 27 July
1971; broken branches and part of main trunk of a large colony, USNM 60348.—
Makapuu, Oahu, 21°18.0’N, 157°32.7'W, 362 m, ‘“‘Sango XIV”’ haul 1, 18 January
1972; 1 small colony lacking holdfast, USNM 60290 (Figs. 18, 20).

VOLUME 95, NUMBER 1 145

TTT

Fig. 18. Fanellia tuberculata (Versluys) from Hawaii, USNM 60290: typical quasi-dichotomous
colony with sterile twig tips. Scale divisions are mm.

Marshall Islands: Bikini Atoll, off Enyu Pass, 11°29’28’N, 165°31'40”E, 212-220
m, dredge sta. 30, 22 August 1947; one branch, broken in two pieces, from a
larger colony, holotype of Caligorgia pseudoflabellum Bayer, USNM 44089 (Figs.
O22).

Japan: Honshu, Ose Zaki S.36°, W 0.8 mi, 65—125 fath., volcanic sand, shells,
rock, Albatross sta. 3716, 11 May 1900; large dichotomously divided branch

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 19. Fanellia tuberculata (Versluys) from Hawaii, USNM 56791: right and left lateral views of
polyp. SEM stereo pairs, x56.

lacking holdfast, USNM 49582 (Fig. 23); smaller branch, part of preceding?,
USNM 57523.—Kyushu, off Kagoshima Wan, 30°54’40’N, 130°37'30"E, 103 -fath.,
stones, Albatross sta. 4936, 16 August 1906; flabellate colony with holdfast, with
broken branches, Caligorgia flabellum det. Nutting, USNM 30029 (Figs. 24, 26);
branch without holdfast, part of preceding?, Caligorgia aspera det. Nutting,
USNM 30075.—Kyushu, Westkuste von Satsuma; small branch of colony det.

VOLUME 95, NUMBER 1 147

Fig. 20. Fanellia tuberculata (Versluys) from Hawaii, USNM 60290: opercular, adaxial marginal,
‘abaxial, and outer-lateral sclerites of polyp. Scale = 0.1 mm.

K. Kinoshita in Zoological Institute, University of Tokyo, USNM 50120.—Kyu-
shu, Kagoshima, 70 fath., 15 July 1899; small branch of colony det. K. Kinoshita
in Zoological Institute, University of Tokyo, USNM 60291 (Fig. 25).
Description.—Branching quasi-dichotomous or loosely pinnate, alternate, com-
pressed or in one plane. Polyps in whorls of 2 or 3 near tips of terminal twigs,
increasing proximally to 5 in some colonies, and as many as 7 or 8 on larger
branches, often becoming irregularly scattered; usually 7 or 8 whorls in 1 cm of
axial length in the terminal twigs, sometimes as few as 6 or as many as 9. Polyps
typically with 6 scales in the abaxial rows, but occasionally as few as 4 and as
many as 8; outer lateral rows with 1 or 2 scales; inner lateral rows with only |
and this may be absent on one or both sides, its place occupied by wide lateral
extension of the outer-lateral scale (OL-1); OL-2 if present may extend laterally
as a curved lobe beneath the inner-lateral marginal, but a separate second inner
lateral (IL-2) has not been observed; 1-3 small scales, commonly 1 or 2, in the
adaxial rows beneath the adaxial operculars. Tentacles without sclerites. Coe-

148 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 21. Fanellia tuberculata (Versluys) from Marshall Islands, USNM 44089, holotype of Cal-
logorgia pseudoflabellum Bayer: top, whorl of polyps, x44; bottom, lateral view of polyp, x56. SEM
stereo pairs.

nenchyme covered with closely fitted, rounded-polygonal plates. All sclerites
sculptured externally by closely placed, serrated, fluted or ridged tubercles
aligned in rows radiating outward from the depositional center (‘‘nucleus’’), in-
ternally by crowded, complex tubercles.

Comparisons.—Kinoshita (1908a:40) remarked upon the similarity of his Cali-
gorgia aspera to C. tuberculata Versluys, but justified its status as a distinct
species on the basis of differences in the form of the external warts of the scales,
and on the absence of the sterile twig tips reported in C. tuberculata by Versluys
(1906:80). These sterile tips are of inconsistent occurrence in Hawaiian material,

Islands, USNM 44089, holotype of Cal-
, abaxial and outer-lateral sclerites of

0.1 mm.

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VOLUME 95, NUMBER 1

polyp. Scale

150 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 23. Fanellia tuberculata (Versluys), quasi-dichotomous colony from Japan, USNM 49582.
Scale in cm.

but it is not possible to determine whether they are present in some colonies but
not in others, because the material was so badly broken during collection. Some
of the specimens with sterile branch tips are infested with colonies of a thecate
hydroid which seem to grow only on these sterile tips, but I cannot determine

VOLUME 95, NUMBER 1 151

Fig. 24. Fanellia tuberculata (Versluys), loose pinnate colony from Japan, USNM 30029. Scale
divisions are mm.

whether the hydroids stimulate their production. Such branch tips are not present
in any of the Japanese specimens examined.

It should be noted that Nutting (1912:60, 61) reported two specimens from one
and the same Albatross station, probably parts of one colony, as two different
species, Caligorgia flabellum (wrongly attributed to Kolliker) and C. aspera Ki-
noshita, the former described as ‘“‘regularly alternate,’’ the latter as ‘‘dividing
dichotomously.”’

Although the holotype colony of Caligorgia pseudoflabellum is pinnately
branched (Bayer 1949:207), the branchlets are widely spaced as well as further
subdivided, and the main axis bends slightly away from the branch at each node,
resulting in a loose pinnate pattern that can easily pass for dichotomous. This is

152 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

‘

i

a;
SY
. . a

ans

s

ey

fae

5
a

PAS
i)

Pay
ap,
—

Eee,
wpsetaec)

‘2
AS
~
\ Af
awe
cw &

Fig. 25. Fanellia tuberculata (Versluys) from Japan; original material of C. aspera determined by
Kinoshita, USNM 60291. Right and left lateral views of polyps. SEM stereo pairs, x56.

VOLUME 95, NUMBER 1 153

eA poe
SON a
SO,

Fig. 26. Fanellia tuberculata (Versluys) from Japan, USNM 30029: opercular, adaxial marginal,
abaxial, and outer-lateral sclerites of polyp. Scale = 0.1 mm.

154 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

clearly seen in the photograph (Bayer 1949: pl. 4, fig. 2), in which the distalmost
part of the right-hand fragment would justifiably be described as dichotomous if
it were broken off from the lower part. As this growth form does not differ
significantly from that illustrated for C. aspera by Kinoshita (1908a: pl. 2, figs. 15,
16), and as Kinoshita observed that ‘‘die junge Kolonie sowohl als der Stam-
mabschnitt der erwachsenen sind beinahe federartig verzweigt’’ it is evident that
this character is unreliable. The distribution of the polyps and the arrangement
of their body sclerites are in such close agreement in C. tuberculata, C. aspera,
C. granulosa, and C. pseudoflabellum that it is probable all four nominal taxa
belong to a single species. Although no authentic material of C. granulosa has
been available to me for examination, the specimen from Albatross sta. 4936 off
Kyushu (Fig. 24) is loosely pinnate and approaches (but does not duplicate) the
close-pinnate form originally described for that species. However, on the strength
of existing data, this specimen must be referred to as C. aspera, leaving C.
granulosa as a distinct nominal taxon pending detailed study.
Distribution.—Sulu Archipelago, Japan, Marshall Islands, Hawaii.

Fanellia corymbosa, new species
Figs. 27-29

Material examined.—One colony, broken in several pieces: Moluccas, off Do-
wotra Island, 0°50’00"S, 128°12'E, 205 fath. (375 m), coral sand; Albatross
sta. D5629, 2 December 1909. Holotype colony, USNM 50146.

Description.—Branching irregularly dichotomous, not in one plane, producing
colonies of corymbose form (Fig. 27). Polyps in pairs, the individuals opposite
or slightly offset, infrequently in whorls of 3 even on the larger branches; polyps
irregularly scattered on the largest branches, absent from the side opposite the
direction in which the branchlets grow; 6 or 7 pairs or whorls in 1 cm of branchlet.
Branches stiff and rather brittle, the terminal branchlets commonly 10 mm long
or less, and 0.75 mm in diameter (exclusive of polyps), with the axis 0.3—0.4 mm
in diameter proximally, tapering to a rather coarse point. Diameter of the largest
main stem about 3.5 mm exclusive of cortex; axis heavily calcified, brittle, surface
inconspicuously grooved longitudinally, appearing nearly smooth, distally creamy
yellow with golden iridescence, proximally brown with bronzy gloss.

Polyps (Fig. 28) with only the 2 abaxial rows of sclerites well developed, 5 or
6 thick plates in each, the boundary between proximal abaxial and cortical plates
difficult to determine; one wide lateral scale in marginal position on both sides,
bearing the outer-lateral and inner-lateral opercular scales; occasionally the lateral
marginal plate is divided into two, thus representing outer-lateral and inner-lateral
plates; one or two (usually one) small adaxial scale beneath each adaxial oper-
cular, but the remaining adaxial surface of the polyps naked. Body plates (Fig.
29) thick, the external tubercles near the distal margin forming blunt dentations
that interlock with the proximal tubercles of the following sclerite, inner surface
covered with fine, complex tubercles; opercular scales triangular, the laterals
only slightly asymmetrical, the adaxials somewhat smaller than the others but
not markedly different in shape; on the inner surface of the operculars a low,
strong longitudinal ridge extends from center to apex, most prominent on the
dominant abaxial, progressively smaller on the outer and inner laterals and adax-

VOLUME 95, NUMBER 1 155

Fig. 27. Fanellia corymbosa, new species; holotype colony, USNM 50146. Width 35 cm.

ials; outer surface convex, a prominent rounded boss at the nucleus, covered
with closely set hemispherical tubercles that become lower and less prominent
toward the edges and elongated in shape toward the apex of the scale. Tentacles
without sclerites.

Coenenchymal sclerites rounded or spheroidal pebbles with hemispherical tu-

156 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Sawn it bi
‘ ot a: %
. oi ee

pak ce

“

4 ~~ S,
* Ei. Ke! Lo). ee. oe
; re
a rae
t ie > Fei
Bn} Bae
LL tke RRR Ngee

of polyp. SEM stereo pairs, x45.

bercles on outer surface, finer complex tubercles on inner surface; largest sclerites
become thick, irregular bodies resembling coarse cobblestone pavement.

Comparisons.—In many ways, this species resembles Kinoshita’s Callogorgia
aspera and C. granulosa, the first definitely and the second possibly a synonym
of Fanellia tuberculata (Versluys), but differs from both in growth form. F.
tuberculata, from the Sulu Archipelago, usually has 2 plates in the outer-lateral
rows and | in the inner-lateral rows, whereas F. corymbosa has only a single
outer-lateral plate on each side, extending into the inner-lateral position where
no separate plate is developed. The lower abaxial plates of corymbosa are much
wider (to 0.6 mm) than those of tuberculata (0.3-0.4 mm). No sterile terminal
branch tips were observed in corymbosa.

VOLUME 95, NUMBER 1

Fig. 29. Fanellia corymbosa, new species; holotype, USNM 50146: opercular, abaxial marginal
and abaxial proximal body sclerites. Scale = 0.1 mm.

158 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Acknowledgments

I am grateful to Dr. P. F. S. Cornelius and Mr. Gordon Paterson of the Coel-
enterate Section, British Museum (Nat. Hist.), who made available for study
paratypes of Hickson’s Caligorgia fraseri. Dr. K. K. Chew supplied, through the
Smithsonian Oceanographic Sorting Center, some of the material of Fanellia
compressa here described. The specimens of Callogorgia formosa and Fanellia
tuberculata from Hawaiian waters were provided by Dr. Richard Grigg. The speci-
mens of C. aspera identified by K. Kinoshita were made available to me in Tokyo
by Dr. K. Takewaki of Tokyo University. All scanning electron micrographs were
made by Mr. Walter R. Brown, chief of the SEM Laboratory, National Museum
of Natural History, Smithsonian Institution. Photographs of colonies were made
by Mr. Michael R. Carpenter, who also prepared the photographic prints re-
produced in this paper. Dr. Stephen D. Cairns kindly reviewed the manuscript.
To all of these I express my thanks.

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Pallas, P. S. 1766. Elenchus zoophytorum sistens generum adumbrationes generaliores et specierum
cognitarum succinctas descriptiones cum selectis auctorum synonymis. Hagae Comitum.
i-xvi + 17-28 + 1-451 pp.

Statius Muller, P. L. 1775. Von den Corallen. Jn Des Ritters Carl von Linne .. . vollstandiges
Natursystem . . . 2(6):i-xvi + 641-960 [1068], pls. 20-37. Nurnberg, Gabriel Nicolaus Raspe.

Studer, T. 1878. Ubersicht der Anthozoa Alcyonaria, welche wahrend der Reise S.M.S. Gazelle um
die Erde Gesammelt wurden.—Monatsbericht der KGniglich Preussischen Akademie der Wis-
senschaften zu Berlin 1878:632-688, pls. 1-5.

. 1887. Versuch eines Systemes der Alcyonaria. Archiv fiir Naturgeschichte 53(1):1-74, pl. 1.

Thomson, J. A., and N. I. Rennet. 1931. Alcyonaria, Madreporaria, and Antipatharia.—Australasian
Antarctic Expedition 1911-14. Scientific Reports (C-Zoology and Botany) 9(3):1—46, pls. 8-14.

Verrill, A. E. 1865. Synopsis of the polyps and corals of the North Pacific Exploring Expedition,

160 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

under Commodore C. Ringgold and Captain John Rodgers, U.S.N., from 1853 to 1856. Col-
lected by Dr. Wm. Stimpson, naturalist to the Expedition. With descriptions of some additional
species from the west coast of North America.—Proceedings of the Essex Institute, Salem,
4:181—196. pls. 5-6.

Versluys, J. 1906. Die Gorgoniden der Siboga Expedition II. Die Primnoidae.—Siboga-Expeditie
Monographie 13a. Pp. 1-187, figs. 1-178, pls. 1-10, chart.

Wright, E. P., and T. Studer. 1889. Report on the Alcyonaria collected by H.M.S. Challenger during
the years 1873-1876.—Voyage of the Challenger, Zoology, 31:i-Ixxii + 1-314, 43 pls.

Department of Invertebrate Zoology, Smithsonian Institution, Washington,
D.C. 20560.

PROC. BIOL. SOC. WASH.
95(1), 1982, pp. 161-166

FRESHWATER TRICLADS (TURBELLARIA) OF NORTH
AMERICA. XIII. PHAGOCATA HAMPTONAE,
NEW SPECIES, FROM NEVADA

Roman Kenk

Abstract.—A new species of the planarian genus Phagocata, P. hamptonae,
is described. It differs from all other North American species of the genus by
having 4 to 11 eyes in the usual position. In the copulatory apparatus, the struc-
ture of the penis deviates from the general plan by having a wide ejaculatory duct
filled with an eosinophilic secretion. The species forms spermatophores that are
inserted into the bursa of the copulating partner.

In an extensive field study of the freshwater triclads of the Lake Tahoe basin
and the surrounding area, Dr. Anne M. Hampton discovered a white planarian
with more than one pair of eyes, that she recognized as being a new taxon. She
kindly sent me a number of live specimens for a more detailed study. Some of
the worms were sexually mature.

Phagocata hamptonae, new species

Type-material.—All type-specimens are deposited in the National Museum of
Natural History, Smithsonian Institution, Washington, D.C. Holotype, set of
sagittal sections on 3 slides (USNM 68009): paratypes. 5 sets of sagittal sections
on 3 slides each (USNM 68010-68014) and one set of transverse sections of
posterior part of body on 3 slides (USNM 68015).

External features (Fig. 1).—The species is unpigmented, white, and at first
glance resembles the eastern Phagocata morgani (Stevens and Boring). Mature
specimens reach a length of 11 mm and a width of 1.3 mm. The anterior end is
truncate, with a straight or somewhat wavy frontal margin (slightly convex in the
center), changing somewhat during gliding locomotion, and with rounded lateral
corners. Behind the anterior end, a slight narrowing or “‘neck’’ may be seen in
the gliding animal. The body soon reaches its maximum width, the lateral margins
running parallel in the greater part of the body length, converging in the last fifth
of the body, and meeting at the bluntly pointed posterior end.

The most characteristic feature of the species is the plurality of the eyes. While
most species of Phagocata have only one pair of eyes or are blind, P. hamptonae
has a small but variable number of eyes (Fig. Ic). Dr. Hampton, who analyzed
37 specimens, found their number to vary between 4 and 11, the mean number
being 6, arranged generally in two longitudinal rows separated by about one-fifth
the transverse diameter of the head. The eyes are rather small, of about equal
size, but fully functional, each equipped with a well-developed pigment cup.

The pharynx measures about 1/5 the body length. its root being inserted anterior
to the middle of the body in mature specimens. The intestine is amply ramified,
which makes it difficult to count the branches of the intestinal trunks. By feeding
some specimens a mixture of beef-liver tissue and carbon powder and later ex-

162 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Phagocata hamptonae: A, Photograph of living specimen after ingestion of a mixture
of beef-liver tissue and carbon powder to show the intestine, x7.5; B, Outline drawing with indica-
tion of eyes, pharynx, and copulatory apparatus; C, Arrangement of eyes, redrawn from sketches
prepared by A. M. Hampton.

amining them under slight compression, the approximate number of branches
could be established: 4—6 heavily ramified branches on either side of the anterior
trunk; the two posterior trunks each bear about 14—19 less ramified lateral branch-
-es and a considerable number of short medial diverticula. Postpharyngeal anas-
tomoses between the two trunks may occur.

Anatomy.—The anterior end shows no special structures (glands or muscles)
that could be interpreted as corresponding to an adhesive organ. The zone of
eosinophilic adhesive glands, that runs along the ventral side of the body margins,
is interrupted in the central area of the frontal margin, where there is a field of
infranucleate ciliated epithelium, apparently a sensory organ. The pharynx has
the normal structure characteristic of the Planartidae, its internal muscular zone
consisting of a thick layer of circular fibers surrounding the pharyngeal canal,
followed by a layer of longitudinal muscles.

The two ovaries or germaria are situated on the medial side of each ventral
nerve cord at the level of the first lateral branches of the intestine. Each ovary
is accompanied by a group of cells about the size of the ovary, situated anterior
to it. Such cell accumulations adjoining the ovaria are widely distributed in tri-

VOLUME 95, NUMBER 1 163

|
|
i
)

| | |
m VEL weKYS de am gD
Fig. 2. Phagocata hamptonae, semidiagrammatic view of copulatory apparatus in sagittal section:

am, male atrium; b, copulatory bursa; bd, bursal duct; bp, penis bulb; de, ejaculatory duct; gp,
gonopore; m, mouth; odc, common oviduct; pp, penis papilla; vd, vas deferens; vs, seminal vesicle.

clads and are usually termed ‘“‘parovaria.’’ In P. hamptonae these cells corre-
spond entirely in size, stainability, and cytoplasmic inclusions (yolk granules) to
the cells of the vitellaria, a fact that confirms the often expressed assumption that
the parovaria have a genetic, if not always functional, relation to the yolk glands.

The numerous testicles are of moderate size, usually measuring less than half
the dorsoventral diameter of the body. They are predominantly ventral and are
arranged in longitudinal rows beginning a short distance posterior to the ovaries
and extending to almost the hind end of the body. The two sperm ducts or vasa
deferentia run along the ventral nerve cords medial to the oviducts. In the region
of the pharynx they expand to form the spermiductal vesicles or ‘‘false seminal
vesicles’ which, filled with sperm, proceed posteriorly to their entrance into the
penis bulb.

The copulatory apparatus (Fig. 2) closely adjoins the pharyngeal pouch. The
- genital atrium is confined to the male atrium (am), since the bursal duct (bd)
proceeds directly to the gonopore (gp) without entering a separate atrial com-
partment. The penis consists of a relatively large bulb (bp) and a conical papilla
(pp). The bulb has its musculature developed principally near its periphery, while
its central part is remarkably devoid of muscle fibers. The lumen of the penis is
clearly divided into two sections. An anterior cavity, the seminal vesicle (vs),
located in the bulb, is lined with an epithelium of cuboidal, apparently apocrine,
cells, the secretions of which must have dissolved during the procedures used in
preparing the sections, so that the cells look empty (see Fig. 3). The posterior
part (de) of the lumen is a rather wide, elongated cavity, separated from the

164 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Phagocata hamptonae, photomicrograph of a sagittal section through the copulatory com-
plex of a specimen that has two spermatophores in the copulatory bursa, <120.

seminal vesicle by a slight constriction and extending from the penis bulb to the
tip of the papilla, tapering gradually in its posterior portion. It is almost com-
pletely filled with strands of a granular, slightly eosinophilic secretion, the origin
of which is not quite clear. It may be in part produced by the epithelial cells
lining the cavity, but traces of apparently the same secretion are seen also in the
parenchymal part of the penis papilla, which would hint at gland ducts entering
from the mesenchyme surrounding the penis bulb. This posterior part of the penis
lumen corresponds by its location to the ejaculatory duct of related species,
although it may have a different function, possibly related to the production of
spermatophores. The two sperm ducts or vasa deferentia (vd) enter the penis
bulb ventrolaterally, turn upward, and open into the seminal vesicle separately.

The two oviducts or ovovitelloducts run along the dorsal side of the ventral
nerve cords up to the level of the male atrium. There they turn dorsally, approach
the midline, and unite in the space between the atrium and the bursal duct. The
common oviduct (odc) thus formed proceeds posteroventrally in a short curve
along the atrial wall and opens into the posterior part of the atrium. The paired
oviducts, after they have left the nerve cords, and the upper part of the common
oviduct receive the highly eosinophilic shell glands.

The copulatory bursa (b) is of variable size and shape. In some of the specimens
it contained remnants of spermatophores (see Fig. 3). Spermatophores were also
visible in some living specimens as glittering spherules shining through the trans-
parent body wall.

The bursal stalk or duct (bd) runs from the bursa posteriorly above the penis
and atrium, then curves ventrally, slightly increasing in diameter, and opens at
the gonopore (gp) to the left of the outlet of the male atrium. No differentiated

VOLUME 95, NUMBER 1 165

end part or vagina is developed and the duct is histologically uniform throughout.
Its musculature consists of a subepithelial layer of circular fibers, surrounded by
a layer of longitudinal muscles.

All epithelia of the copulatory complex are nucleate, including the common
oviduct, while the paired oviducts show an infranucleate epithelium.

Distribution and ecology.—Phagocata hamptonae was collected by Dr. Anne
M. Hampton in an irrigation ditch 400 m east of the ranch house on the Gene
Scossa Ranch, Gardnerville, Douglas County, Nevada, on 5 April 1981 (water
temperature, 15°C, pH 7.5) and 5 July 1981. Dr. Hampton sent me samples from
both collections, containing some sexually mature and some immature specimens.
The worms were placed in cultures kept at 14°C. They readily accepted beef liver
as food.

Taxonomic position.—The genus Phagocata is admittedly a very heteroge-
neous genus, distributed widely in the Northern Hemisphere in both Eurasia and
North America (see also discussion by Ball and Gourbault 1975:11-13). The mem-
bers of the genus conform in the basic structure of the copulatory apparatus,
though they may differ in many other characters that are of taxonomic signifi-
cance. In some species the testicular zone extends to the tail end, in others only
to the level of the mouth; some are pigmented, others white; they generally have
one pair of eyes, but some are blind, and the present species has multiple eyes
(as does P. uenoi Okugawa from Manchukuo, see Okugawa 1939:157). It must
be stressed that the eyes are completely formed eyes with no intergrades between
functional eyes and apparently nonfunctional pigment clumps, such as are seen
in P. morgani polycelis Kenk (1935:103), Dendrocoelopsis americana (Hyman)
(Kenk 1973:14) and some species of Dendrocoelum. The structure of the penis
may vary in the genus within wide limits even in species that are closely related
both morphologically and zoogeographically. Within certain geographic regions
we may find groups of species that are undoubtedly monophyletic and could be
gathered into separate subgenera or even genera. Phagocata hamptonae has the
zone of testes approaching the posterior end of the body, a character it shares
with nine of the North American species of the genus. On the other hand, P.
morgani (Stevens and Boring), P. bursaperforata Darlington, P. bulbosa Kenk,
P. angusta Kenk, and P. holleri Kenk have no testes behind the copulatory
apparatus. Phagocata hamptonae is monopharyngeal as are all the other North
American species except P. gracilis (Haldeman), P. woodworthi Hyman, and P.
nordeni Kenk. The new species may well deserve to be placed in a separate
subgenus, principally on account of the aberrant morphology of its penis. This
should be done, however, within the framework of a reexamination of the entire
genus.

The species is named in honor of its collector, Dr. Anne Marie Hampton of
Tahoe Paradise, California.

Acknowledgments

Thanks are due to Dr. Hampton for the transmittal of the material and for her
generous collaboration in the study of the new species. Dr. John C. Harshbarger
of the Smithsonian Institution was helpful in the preparation of the photomicro-
graph and Dr. Marian H. Pettibone kindly reviewed the manuscript.

166 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Literature Cited

Ball, I. R., and N. Gourbault. 1975. The morphology, karyology, and taxonomy of a new freshwater
planarian of the genus Phagocata from California (Platyhelminthes: Turbellaria).—Life Sci-
ences Contributions, Royal Ontario Museum 105:1-19.

Kenk, R. 1935. Studies on Virginian triclads.—Journal of the Elisha Mitchell Scientific Society

51:79-133.

. 1973. Freshwater triclads (Turbellaria) of North America. VI. The genus Dendrocoelopsis.—

Smithsonian Contributions to Zoology 138:1-16.

Okugawa, K. 1939. Probursalia (Triclada-Paludicola) of Manchoukuo.—Annotationes Zoologicae Ja-
ponenses 18:155-165.

Department of Invertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
95(1), 1982, pp. 167-187

CLASSIFICATORY REVISIONS IN GAMMARIDEAN
AMPHIPODA (CRUSTACEA), PART 2

J. Laurens Barnard and Gordan S. Karaman

Abstract.—The new family Paracalliopidae is established and the family Pseud-
amphilochidae is recognized from provisional proposal.

New genera are described as follows: Relictomoera for Paramoera relicta and
Sternomoera for P. yezoensis and P. japonica; Nasageneia for Pontogeneia
nasa; Aurohornellia for Tulearogammarus sinuatus; Lupimaera for Maera lu-
pana; Tegano for Melita seticornis; Maleriopa for Eriopisella dentifera; Texi-
weckeliopsis for Texiweckelia insolita; Holsingerius for T. samacos; Indocalliope
for Paracalliope indica; Feriharpinia for Harpinia ferenteria; Torridoharpinia
for Proharpinia hurleyi.

New synonymies are provided for Afrochiltonia (=Austrochiltonia), Cerado-
copsis (=Maeracunha), Hornellia (=Metaceradocus). Echiuropus 1s raised to full
generic level. New lists of species are provided for Pseudomoera, Paracalliope
and Proharpinia.

More ‘‘armchair’’ revisions in Amphipoda must be presented before we can
issue our forthcoming ‘‘The Families and Genera of Gammaridean Amphipoda,”’
a sequel to that done by J. L. Barnard (1969b). This time, freshwater taxa will
be included with the marine in the same treatise.

Our first part is cited as Karaman and Barnard (1979).

Ceinidae
Afrochiltonia K. H. Barnard, new synonymy

Afrochiltonia K. H. Barnard, 1955:93 (Chiltonia capensis K. H. Barnard, 1916,
original designation).

Austrochiltonia Hurley, 1958:767 (Hyalella australis Sayce, 1901, original des-
ignation).

Griffiths (1976) made a case for synonymizing these two genera but mistakenly
assumed Austrochiltonia took priority. It does not and therefore the situation
should appear as shown above.

Eusiridae
Pseudomoera Schellenberg, revised

Pseudomoera Schellenberg, 1929:281 (Atyloides gabrieli Sayce, 1901, monotypy).

Until now this genus has been monotypic but we transfer into this genus Aty-
loides fontana Sayce (1902) which for years has been placed in Paramoera. The
type-species is characterized by the loss of medial setae on the inner plate of
maxilla | whereas P. fontana has full setal armament on that plate like Para-
moera. However, Pseudomoera can now be distinguished from Paramoera on
the presence of geniculate lobes on the wrists of the gnathopods and this action

168 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

removes from Paramoera a clearly alien species and places it precisely contig-
uous to its apomorph. This creates an unusual genus in which maxillary setae are
variable but this is perhaps preferable to adding yet another monotypic genus to
the pool.

Species.—fontana (Sayce, 1902a); gabrieli (Sayce, 1901); Australia, Victoria,
streams between 457 and 915 m of altitude.

Relictomoera, new genus

Type-species.—Paramoera relicta Ueno, 197la.

Body slender, compressed, smooth, urosomites free. Rostrum obsolescent;
lateral cephalic lobes sinusoid; with notch in middle, anteroventral margin of
head weakly produced. Eyes round, tiny, or vestigial, or absent.

Antennae elongate, | longer than 2, peduncular articles of antenna | progres-
sively shorter (ratio 21:19:12), article | as long as or longer than head, articles
2—3 scarcely shorter than article 1; article 3 not produced; article 1 of primary
flagellum ordinary, accessory flagellum 1-articulate, barrel shaped.

Labrum [?entire, ?subrounded, broader than long]; epistome unproduced. Mo-
lar triturative, columnar, article 2 of palp poorly lobed, article 3 shorter than 2
(ratio = 4:16:13). Labium: inner lobes [?absent].

Maxilla 1: inner plate with 5 apical setae, outer plate with 9 spines, palp long,
article 1 short. Maxilla 2: inner plate not broader nor longer than outer plate,
narrow, inner plate with facial row of 4 setae but no other medial setae. Maxil-
liped: inner plate relatively long, outer plate slightly shorter than inner, not spi-
nose medially; palp of 4 articles, 4 slightly shorter than 3, 3 unlobed, 4 not spinose
along inferior margin, unguiform, with nail.

Coxae short but contiguous, almost glabrous, coxa | not produced anteriorly
nor expanded ventrally, coxa 4 with weak posterior lobe, excavate.

Gnathopods feeble, slender, diverse, 2 larger than |, subchelate, not eusirid,
medium, wrist of both as long as hand, without posterior lobe, with few short
posterior setae, propodi rectangular or weakly expanded, gnathopod 2 especially.

Pereopods 3-7 ordinary, simple, dactyls simple, article 2 not anteriorly lobate,
on pereopods 5—7 weakly expanded, weakly or moderately lobate.

Epimeron 3 smooth. Pleopods with peduncle dominating rami.

Outer rami of uropods 1-2 slightly shortened or not; rami with lateral and
dorsal spines. Uropod 3 ordinary, scarcely extended beyond uropod |, peduncle
without large process, rami lanceolate, aequiramous, |-articulate.

Telson ordinary, cleft two thirds, apices with short or thin apical armaments.

Coxal gills present on somites 2-6, ovate, some pediculate. Oostegites broad.
Sternal gills absent.

Variables.—Telson elongate (tsushimana).

Relationship.—Like Paramoera but head with unusually sinusoid anterolateral
margin. See Sternomoera. Differing from Awacaris Ueno by the normal palp of
maxilla 1, the normal head and the stronger hand and palm of gnathopod 1.
Differing from Apherusa in the stronger sinusoid cephalic lobes, the deeply cleft
telson, and the presence of an accessory flagellum.

Species.—relicta (Ueno, 197la); tsushimana (Ueno, 1971b); Goto and Tsu-
shimana Islands south and west of Kyushu, Japan, hypogean.

VOLUME 95, NUMBER 1 169

Etymology.—Name composed of root from species name plus ““moera’’ a com-
mon root of genera in Eusiridae. Feminine.

Sternomoera, new genus

Type-species.—Paramoera yezoensis Ueno, 1933.

Body ordinary, compressed, smooth, urosomites free. Rostrum very small,
lateral cephalic lobes rounded or mammilliform, anteroventral margin of head
scarcely produced, with normal sinus. Eyes reniform.

Antennae elongate, | longer than 2, peduncular articles progressively shorter
(ratio = 30:18:12), article 1 shorter than head, article 3 not produced; article | of
primary flagellum short, basal articles proliferate, accessory flagellum 1-articu-
late, scale-like.

Labrum entire, subrounded, as long as broad; epistome unproduced. Molar
triturative, columnar, article 2 of palp unlobed, article 3 as long as 2 (ratio =
4:9:10). Labium: inner lobes absent.

Maxilla 1: inner lobe with many medial setae, outer plate with 7 spines, palp
long, article 1 short. Maxilla 2: inner plate not broader nor longer than outer,
inner plate with scarcely submarginal facial row of many setae and several other
medial setae. Maxilliped: inner plate not relatively long, outer plate slightly short-
er than inner, medially spinose, palp of 4 articles, 4 shorter than 3, 3 unlobed, 4
not spinose along inferior margin, weakly unguiform, [?with nail].

_Coxae ordinary, poorly setose, coxa | not produced anteriorly or expanded
ventrally, coxa 4 with posterior lobe, excavate.

Gnathopods alike in female, almost feeble, subchelate, not eusirid, small wrist
of both scarcely shorter than hand, without posterior lobe, with numerous pos-
terior setae, hands rectangular; male gnathopods larger, 2 larger than 1, wrists
broadly lobate, hands ovate, posterior margins swollen, palms oblique, with few
large clavate or peg spines.

Pereopods 3-7 ordinary, simple, dactyls simple; article 2 of pereopods 3-4 not
anteriorly lobate, of pereopods 5-7 poorly expanded, lobate, poorly armed; or of
pereopod 6 slightly narrowed. Pleopods [?ordinary]. Epimeron 3 serrate sparsely
and minutely.

Outer rami of uropods 1-2 not or scarcely shortened; rami with lateral and
dorsal spines. Uropod 3 ordinary, not extended beyond uropod 1, peduncle with-
out large process, rami lanceolate.

Telson ordinary, cleft, apices with small apical armaments. Coxal gills [2-7],
ovate. Oostegites broad. Thorax with sternal gills.

Variables.—Apices of telson poorly (yezoensis) to densely setose (japonica)
apically.

Relationship.—Like Paramoera but sternobranchiate. Not of modern crango-
nyctid affinity because of precisely aequiramous uropod 3, short peduncle of
antenna 1, basally proliferate flagellum of antenna 1. Not bogidiellid because of
the general gnathopodal facies and presence of sternal gills. Differing from Eo-
niphargus in the magniramous, aequiramous uropod 3 and vestigial accessory
flagellum. Differing from Relictomoera in the presence of sternal gills and the
normal head.

170 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Species.—hayamenensis (Stephensen, 1944); japonica (Tattersall, 1922); ye-
zoensis (Ueno, 1933); Japan, streams.

Etymology.—Name composed of “‘sterno’’ referring to sternal gills and
‘‘moera’’ a root used commonly for genera in Eusiridae. Feminine. 3

Nasageneia, new genus

Type-species.—Pontogeneia nasa J. L. Barnard, 1969c.

Body slender, compressed, smooth. Rostrum large, lateral cephalic lobes or-
dinary, anteroventral margin of head scarcely produced. Eyes reniform.

Antennae subequal, peduncular articles of antenna | progressively shorter,
article 1 shorter than head, article 3 weakly produced; article 1 of primary fla-
gellum ordinary to short, accessory flagellum absent.

Labrum entire, subrounded, broader than long, epistome unproduced. Molar
triturative, columnar, article 2 of palp unlobed, article 3 shorter than 2. Labium:
inner lobes absent.

Maxilla 1: inner lobe with medial and 2 apical setae, palp long, article 1 short.
Maxilla 2: inner plate not broader but slightly longer than outer, inner plate
without facial row of setae but with other medial setae, few, large, at least one
slightly submarginal. Maxilliped: inner plate not relatively long, outer plate slight-
ly shorter than inner; palp of 4 articles, 4 slightly shorter than 3, 3 unlobed, 4 not
Spinose along inferior margin.

Coxae ordinary to short, coxa | not produced anteriorly or expanded ventrally,
coxa 4 without posterior lobe, excavate.

Gnathopods diverse, of similar size, subchelate, not eusirid, medium, wrist of
both shorter than hand, only gnathopod 2 with strong posterior lobe extending
distad, wrist without numerous long posterior setae, hands rectangular in female,
inflated in male, in latter with posterior spines outside limits of oblique palm.

Pereopods 3-7 ordinary, simple, dactyls simple, article 2 not anteriorly lobate.

Epimeron 3 serrate.

Outer rami of uropods 1-2 shortened; rami with lateral and dorsal spines.
Uropod 3 ordinary, not extended beyond uropod 1, peduncle with small process,
rami lanceolate.

Telson ordinary, weakly cleft, apices without long apical armaments.

Relationship.—Like Tethygeneia but epimeron 3 serrate and hands of male
gnathopods with posterior spines well outside palmar limits as in Gondogeneia.
Calceoli tending to be much more strongly anthurial than in Tethygeneia, with
one lobe quite linguiform.

Species.—nasa (J. L. Barnard, 1969c, 1979); quinsana J. L. Barnard, 1964,
1969a, 1979); marine, warm temperate California and Mexico, 0-1 m.

Etymology.—Name composed of “‘nasa’’ from species name plus “‘geneia’’ a
name fragment commonly encountered in Eusiridae. Feminine.

Paramoera stephenseni Barnard and Karaman, new name

Paramoera brachyura Stephensen, 1949:18, figure 6 [homonym to Paramoera
brachyura Schellenberg, 1931:201-202, figure 102].

Etymology.—Named for K. Stephensen.

VOLUME 95, NUMBER 1 171

Gammaridae
Aurohornellia, new genus

Type-species.—Tulearogammarus sinuatus Ledoyer, 1967a.

Body [?ordinary], pleon [?and urosome] dorsally crenulate transversely on pos-
terior segmental margins, each of pleosomite 2 to urosomite 2 with 2 dorsal
spines, [?urosomites free]. Rostrum medium, lateral cephalic lobe sharply con-
ical, sinus present.

Antennae elongate, antenna | shorter than 2, ratio of peduncular articles =
23:25:10, primary flagellum longer than peduncle, accessory flagellum 2-articu-
late. Antenna 2 slender.

Labrum broader than long, entire, rounded. Mandibular incisor toothed, molar
triturative, ratio of palp articles = 4:17:15, article 3 linear, stubby, setae = ADE.
Inner lobes of labium present. Maxillae moderately setose medially, inner plate
of maxilla 1 ovate, very short, moderately setose apicomedially, outer plate with
[?7] spines, palps 2-articulate, [?symmetrical]. Inner plate of maxilla 2 [?with
oblique facial row of setae]. Outer plate of maxilliped [?spinose medially, article
3 of palp unlobed, dactyl shorter than 3, unguiform, with nail].

Coxae long, with long setae, coxa | slightly expanded apically, coxa 3 much
smaller than | or 2, coxa 4 long but unlobed. Gnathopods feeble, slender, almost
simple, wrists elongate, hands elongate, thin, palms scarcely evident, gnathopod
2 thinner than gnathopod 1 (male not clarified).

Pereopods 3-4 ordinary. Article 2 of pereopods 5—7 expanded, moderately to
strongly lobate, serratosetulose posteriorly; pereopod 7 elongate, dactyl
[?elongate and setose].

Pleopods [?ordinary]. Rami of uropods 1-2 marginally spinose, outer rami
slightly shortened, peduncle of uropod | [?with basofacial spine]. Uropod 3 slight-
ly extended, magni- or variramous, rami thin, outer ramus with long article 2,
inner ramus reaching apex of article 1 on outer ramus. Telson of ordinary length,
fully cleft, lobes tapering, poorly armed but with dorsal spinules.

Coxal gills [?2—7, ovate]. Oostegites [?narrow].

Relationship.—Standing between Hornellia and Megaluropus; like Hornellia
but coxa 3 reduced and like Megaluropus but rami of uropod 3 thin.

Species.—sinuata (Ledoyer, 1967a); Madagascar, shallow water.

Etymology.—From “‘aurora,’’ dawn, and ‘‘Hornellia,’’ a related genus. Fem-
inine.

Ceradocopsis Schellenberg, new synonymy

Ceradocopsis Schellenberg, 1926:364 (Ceradocopsis kergueleni Schellenberg,
1926, monotypy).

Maeracunha Stephensen, 1949:22 (Maeracunha tristanensis Stephensen, 1949,
monotypy) [new synonym].

Lateral cephalic lobes broadly rounded, sinus present but mandible inserted
there, not antenna 2.

Antennae elongate, antenna | longer than 2, ratio of peduncular articles =
14:12:6, flagellar ratio = 20:7, accessory flagellum with 4 articles. Antenna 2
slender.

IZ PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Mandibular incisor almost smooth, molar small and poorly triturative, palp
small, ratio of articles = 6:15:12, article 3 linear, setae = E and sparse. Inner
lobes of labium absent. Maxillae medially setose, inner plate of maxilla 1 pyriform
to triangular, with medial setae on apical half only, outer plate with 7-11 spines,
palps [?symmetrical]. Inner plate of maxilla 2 with oblique row of facial setae.
Outer plate with naked medial margin, palp article 3 weakly lobate, dactyl shorter
than 3, unguiform, lacking or bearing nail.

Coxae of medium extension, poorly setose, coxa 1 expanded below, coxa 2
with anteroventral acuity, coxae 3-4 weakly shorter than 1-2, coxa 4 unexcavate
posteriorly, coxa 5 as long as 4. Gnathopods diverse, gnathopod | small, of
Melitid form, wrist scarcely elongate, scarcely lobate, hand longer, rectangular,
palm weakly oblique, short, gnathopod 2 enlarged, wrist short to medium, lobate
or weakly, hand elongate, large, rectangular, palm weakly oblique, weakly sculp-
tured or strongly spinose.

Article 2 of pereopods 5-7 expanded, alike, lobate, poorly setose, serrate and
straight posteriorly, appendages otherwise stout.

Rami of uropods 1-2 subequally extended, marginally spinose, peduncle of
uropod | with basofacial spine. Uropod 3 not extended, short, magniramous,
dispariramous, outer ramus with small article 2. Telson of ordinary length, fully
cleft, lobes tapering, moderately to strongly armed apically.

Coxal gills [?2—-6]. Oostegites [?slender].

Variants.—Outer plate of maxilla 1 with I1 spines (peke) but other species
poorly known; though uropod 3 stated as magniramous, actually appearing to be
parviramous kind with reduced outer ramus now so small as to match inner.

Relationship.—Differing from Maera in the strongly setose maxilla 2 (facial)
and the strong article 2 on the outer ramus of uropod 3. Differing from Ceradocus
by uropod 3 in the same way stated for Maera. Differing from Paraceradocus in
the short dispariramous uropod 3 and short, slender antenna 2. Differing from
Ceradocoides in the short, dispariramous uropod 3 and fully cleft telson. Differing
from Ceradocus in the short dispariramous uropod 3. Differing from various
genera near Melita in the absence of inner lobes on the labium and the miniatur-
ized uropod 3.

Species.—kergueleni Schellenberg, 1926 (Bellan-Santini and Ledoyer 1974);
peke J. L. Barnard, 1972b; tristanensis (Stephensen, 1949); Antarctica and an-
tiboreal, especially insular.

Echiuropus Sowinsky, new synonymy

Echiuropus Sowinsky, 1915:55 (Echiuropus macronychus Sowinsky, 1915, mono-
typy).

[Asprogammarus Bazikalova, 1975:38 (no type-species, therefore unavailable)].

[Smaragdogammarus Bazikalova, 1975:64 (no type-species, therefore unavail-
able)].

Bazikalova’s new taxa belong with Echiuropus; she reduced Echiuropus to
subgeneric level under Asprogammarus which is illegal under ICZN rules. Until
type-species are selected by Bazikalova the two new taxa are unavailable. We
trust biologists will allow Bazikalova to rectify this problem rather than capturing
the taxa for themselves.

VOLUME 95, NUMBER 1 173

Hornellia Walker, new synonymy

Hornellia Walker, 1904:268 (Hornellia incerta Walker, 1904, monotypy).—Le-
dover, 1973:29:

Metaceradocus Chevreux, 1925:304 (Metaceradocus perdentatus Chevreux,
1925, monotypy) [new synonym].

Tulearogammarus Ledoyer, 1967:129 (Tulearogammarus peresi Ledoyer, 1967,
original designation, = Hornellia incerta Walker).

Body ordinary, pleon and urosome dorsally crenulate transversely on posterior
segmental margins, urosomites free, often with dorsal articulate spines. Rostrum
small, lateral cephalic lobes rounded-quadrate, sinus present. Eyes present.

Antennae elongate, extending subequally or antenna 2 longer, ratio of pedun-
cular articles on antenna | = 20:13:7, primary flagellum longer than peduncle,
accessory flagellum 2—5 articulate. Antenna 2 ordinary.

Labrum broader than long, notched (type) or entire. Mandibular incisor
toothed, molar triturative, ratio of palp articles = 4:11:10 or 4:18:15, article 3
linear (type) or falcate, setae = (AB)DE. Inner lobes of labium present. Inner
plate of maxilla | ovate, apically and part to all medially setose, outer plate with
11 spines, palps [?symmetrical]. Inner plate of maxilla 2 with oblique facial row
of setae. Outer plate of maxilliped medially spinose, palp article 3 unlobed, dactyl
{?shorter than 3, with nail].

Coxae ordinary, poorly setose, coxa | apically expanded, coxa 4 lobate.
Gnathopods feeble, slender, scarcely subchelate, wrists elongate, lobed (type) or
unlobed, hands ovate to rectangular, hand of gnathopod 2 elongate, palms
oblique, weak, poorly defined.

Pereopods 3-4 ordinary. Article 2 of pereopods 5-7 slightly expanded, poorly
setose, scarcely lobate.

Pleopods ordinary, rami of uropods 1-2 marginally spinose, only outer of uro-
pod 2 shortened, uropod | with basofacial spine [in several species, type
unknown]. Uropod 3 extended, magniramous, aequiramous, peduncle slightly
elongate, rami elongate, lanceolate, setose, outer 1-2 articulate. Telson elongate,
deeply cleft, lobes tapering, notched (type), weakly spinose apically and medially
and dorsally (type) and laterally.

Coxal gills 2-7, ovate. Oostegites slightly broadened to slender.

Variants.—Uropod 3 without article 2 on outer ramus (perdentata, occiden-
talis, micramphopus, vesentiniae), telson fully cleft (occidentalis, micrampho-
pus): gnathopod 2 very linear (micramphopus); Metaceradocus definable as sub-
genus with micramphopus, occidentalis, perdentata and vesentiniae: telson fully
cleft, article 2 of outer ramus on uropod 3 absent.

Relationship.—Differing from Elasmopoides and Maeropsis and Ceradocus in
the feeble gnathopod 2 of both sexes; somewhat ancestral to Maerella and Jer-
barnia but anterior coxae normal, not reduced or diversified and maxillipedal
dactyl well developed; differing from the Eriopisella group in the magniramous
uropod 3 and well setose maxillae.

Species.—incerta Walker, 1904 (= peresi Ledoyer, 1967) (Ruffo 1969); mi-
cramphopus (Stebbing, 1910); occidentalis (J. L. Barnard, 1959); perdentata
(Chevreux, 1925); vesentinae (Ruffo, 1969); whakatane (J. L. Barnard, 1972b);
marine, warm-temperate and tropics, shallow water.

174 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Lupimaera, new genus

Type-species.—Maera lupana J. L. Barnard, 1969a.

Body slender, urosomites free, naked. Rostrum obsolescent, lateral cephalic
lobes mammilliform. Antennae medium to short, antenna 1 scarcely longer than
2, ratio of peduncular articles = 16:8:3, primary flagellum not longer than pedun-
cular article 1, with 5 articles, accessory flagellum 3-articulate, more than half as
long as primary flagellum. Antenna 2 also short, flagellum 4-articulate, not longer
than article 5 of peduncle. Ratio of mandibular palp articles = 2:7:5, article 3
linear, setae = DE. Inner lobes of labium present. Maxillae not setose medially,
inner plate of maxilla 1 rectilinear, with 3 apical setae, outer plate with 9 spines,
palps symmetrical. Inner plate of maxilla 2 with 2 medial and 1 facial setae.
Outer plate of maxilliped medially spinose, palp article 3 unlobed, dactyl shorter
than 3, very short, unguiform, with nail.

Coxae of ordinary dimensions, poorly setose, coxa 1 weakly expanded and
lobed anteroventrally, coxa 4 not lobate, coxa 5 as long as 4. Gnathopods diverse,
not sexually dimorphic, gnathopod | small, wrist elongate, unlobed, hand sub-
rectangular, palm short, slightly oblique; gnathopod 2 enlarged, wrist short,
lobed, hand large, subrectangular, palm oblique, short, sculptured.

Pereopods 3-4 ordinary. Article 2 of pereopods 5-7 weakly expanded, weakly
lobate, posterior margins poorly setose, weakly convex; article 2 of pereopod 5
not longer than coxa 5 (distinction from Maera); pereopods short.

Outer rami of uropods 1-2 slightly shortened, all rami marginally densely spi-
nose, spines small, peduncle of uropod 1 with 1-2 basofacial spines. Uropod 3
not extended, very short, magniramous, almost aequiramous, rami short, scarcely
longer than peduncle, spinose, spines short (distinction from Maera). Telson
short, almost fully cleft, lobes tapering, apices weakly spinose, notched.

Coxal gills 2-6, ovate. Oostegites [?narrow].

Relationship.—Differing from Maera, Paraweckelia, and Meximaera in the
small article 2 of pereopods 5-7, article 2 of pereopod 5 not being longer than
coxa 5; rami of uropods I-—3 with spines all shortened; article 2 of antenna | only
half as long as article 1. More than 55 species of Maera do not conform to this
group of distinguishing characters noted for Lupimaera. Distinguished from Ce-
radocus, which it closely resembles, in the poorly setose maxillae, short uropod
3 and short article 2 of antenna 1; from Ceradomoera in the symmetrical gnath-
opod 2 and smooth pleon; from Maeropsis in the poorly setose maxilla 2; from
Tfalukia in the normally long article 2 of mandibular palp; from Paraceradocus
in the poorly setose maxilla 2 and linear mandibular palp; from Anelasmopus by
the poorly setose maxillae; and from Ceradocoides in the deeply cleft telson and
poorly setose maxillae.

Species.—lupana (J. L. Barnard, 1969a); California, intertidal.

Etymology.—Name composed of “‘lupana’’ and “‘Maera,’’ a related genus.
Feminine.

Lupimaera lupana (J. L. Barnard), new combination
Fig. 1

Maera lupana J. L. Barnard, 1969a:122, figure 20.

Added description.—Right and ieft incisors each with 3 teeth; right lacinia
mobilis bifid and complex (see illustration), left 5-dentate; right rakers 6, left 7;

VOLUME 95, NUMBER 1 175

D

Fig. 1. Lupimaera lupana (J. L. Barnard), male ‘‘h’’ 3.95 mm: A, Lower lip; B, Maxilla 2; C,
Maxilla 1; D, Left incisor and lacinia mobilis; E, Maxilliped; F, Right mandible.

molars of medium size, weakly triturative, each with long seta; palps alike on
either side, article 1 short, article 2 with 3 inner subbasal setae, article 3 shorter
than 2, weakly curved but not considered falciform, of linear form but short,
setae = D2E3. Lower lip with fleshy inner lobes bearing sublobes; outer lobes
normal. Inner plate of maxilla 1 bluntly subconical, apex with 2-3 setae; outer
plate with 9 normal spines; palps symmetrical, 2-articulate; article | slightly elon-
gate, article 2 truncate, with 6 apical setae and 2 medial setules. Plates of maxilla
2 extending equally, outer broader than inner, latter with 2 thin medial setae, one
facial seta near apex assumed to represent facial row. Plates of maxillipeds normal
(illustrated), though article 2 elongate, palp smaller than usual in family, dactyl
short, scarcely unguiform, with large nail.

Coxal gills on segments 2-6 ovate. Oostegites unknown.

Material.—J. L. Barnard Station 41, California, Goleta, 6 July 1961, 3 m, rhi-
zomes of Macrocystis pyrifera, male ‘*h’’ 3.95 mm (mouthparts illustrated).

Remarks.—Now that this species is segregated in its own genus the mouthparts
are herein illustrated; they are not significantly distinct from those of Maera as

176 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

originally implied. The generic distinctions from Maera lie in the posterior per-
eopods (see above).
Distribution.—California, Goleta, in Macrocystis rhizomes, 3 m.

Maleriopa, new genus

Type-species.—Eriopisella dentifera Ledoyer, 1978.

Body slender, urosomites [?free, naked]. Rostrum small, lateral cephalic lobes
weak, rounded, sinus absent. Eyes present.

Antennae moderately extended, ratio of peduncular articles = 26:19:8, primary
flagellum slightly longer than peduncle, accessory flagellum 2-articulate. Antenna
2 ordinary.

Labrum [?entire, rounded]. Mandibular incisor toothed, right mandible with 3
rakers, molar weakly triturative, ratio of palp articles = 3:10:9, article 3 linear,
setae = E, sparse. Inner lobes of labium present, fleshy, mandibular lobes ex-
tended. Maxillae not setose medially, inner plate of maxilla | subrectangular,
with 2 distal setae, outer plate with about 7 spines, palps 2-articulate,
[?symmetrical]. Inner plate of maxilla 2 without facial and medial setae. Outer
plate of maxilliped [?medially spinose, article 3 of palp unlobed, dactyl shorter
than 3, unguiform, with nail].

Coxae of ordinary length, poorly setose, coxa | tapering, coxa 4 [?unlobed].
Gnathopods small to moderate, gnathopod | with elongate wrist, hand subrect-
angular, palm weakly oblique; gnathopod 2 moderately enlarged, wrist short,
strongly lobed, lobe bending distad, hand subrectangular, weakly expanded api-
cally, palm oblique, well defined, strongly spinose.

Pereopods 3-4 ordinary. Article 2 of pereopods 5—6 weakly expanded, mod-
erately lobate, posterior margin straight, of pereopod 7 expanded, lobate, pos-
terior margin convex, all posterior margins weakly setose; dactyls ordinary.

Pleopods [?ordinary]. Rami of uropods 1-2 extending equally, marginally spi-
nose except for outer ramus of uropod 1, latter [?with basofacial spine]. Uropod
3 slightly extended, parviramous, article 2 on outer ramus short. Telson short,
deeply cleft, lobes broad but weakly tapering, each apex with subapical spine.

Coxal gills [?2-6, ovate]. Oostegites [?narrow].

Relationship.—Differing from Eriopisa, Victoriopisa and Psammogammarus
in the loss of medial setae on the maxillae; from Jegano in the presence of article
3 on the mandibular palp, the short article 2 of antenna | and the loss of medial
setae on maxilla 2; from Eriopisella in the slightly enlarged gnathopod 2 with
strong, spinose palm.

The type-species is assumed to have a direct origin distinct from Paraniphargus
as reflected in the tapering coxa 1, presence of eyes, large lobe on wrist of
gnathopod 2, more strongly lobate and larger article 2 of pereopods 5—7, presence
of article 2 on the outer ramus of uropod 3 and the loss of marginal spines on the
outer ramus of uropod 3.

Species.—dentifera (Ledoyer, 1978); Mauritius, sublittoral, 1 species.

Etymology.—Name contrived from parts of ‘‘Mauritius’’ and “‘Eriopisa”’’ with
an ‘“‘ell’’ thrown in between for euphony. Feminine.

Tegano, new genus

Type-species.—Melita seticornis Bousfield, 1970.

VOLUME 95, NUMBER 1 177

Body smooth, urosomite 2 with small middorsal posterior mucronation. Ros-
trum obsolescent, lateral cephalic lobes strongly mammilliform.

Antennae elongate, slightly thickened, antenna | scarcely longer than 2, ratio
of peduncular articles = 12:15:5, ratio of flagella = 30:3, accessory flagellum 2-
articulate. Antenna 2 gland cone large.

Ratio of mandibular palp articles = 4:8:0 or 3:10:0 (article 3 absent), apical
setae of article 2 = E (one long only). Inner lobes of labium well developed,
fleshy. Maxillae moderately setose medially, inner plate of maxilla | ovate, with
6 apical and medial setae, outer plate with ‘‘9-10”’ (but see figure) spines, palps
[?asymmetric]. Inner plate of maxilla 2 with several marginal medial setae, non-
facial setae. Outer plate of maxilliped minutely spinose medially, dactyl with tiny
nail.

Coxae of ordinary length, poorly setose, coxa | undilate, coxa 4 unlobed.
Gnathopods 1-2 diverse, gnathopod | the smaller, of Melitid form, wrist weakly
elongate, hand subrectangular or trapezoidally expanded apically, palm trans-
verse, convex in female, acquiring rugose process in terminal males, gnathopod
2 slightly enlarged, wrist of medium length, scarcely lobed (broadly), hand elon-
gate, subrectangular in female, with oblique short palm, in male larger, more
ovate, palm longer, softly excavate, dactyl longer and more curved.

Article 2 of pereopods 5-7 scarcely expanded, scarcely lobate or not postero-
ventrally, poorly setose (only setulose).

Rami of uropods 1—2 extending equally, marginally spinose, peduncle of uropod
1 with basofacial spine. Uropod 3 extended, parviramous, outer ramus elongate,
with medium article 2. Telson short, cleft to base, lobes leaf-like, tapering api-
cally, sparsely setose apically.

Coxal gills 2-6 ovate, that on pereonite 2 pediculate. Oostegites narrow.

Description.—Antennae of male with whorls of setae on peduncles and flagella.

Relationship.—Dijffering from Melita in the reduced mandibular palp.

Species.—seticornis (Bousfield, 1970); Solomon Islands (Rennell) and Bis-
marck Archipelago (Mussau), anchialine.

Etymology.—Named for Lake Tegano, Rennell Island, type-locality of the
type-species. Masculine.

Hadziids and Weckeliids

A Weckeliid lacks eyes and has aequiramous uropod 3. A Hadziid lacks om-
matidial eyes though occasionally has ocular pigment but uropod 3 is disparira-
mous. Hadziids further differ from neighboring blind taxa in the elongate uropod
3, cleft telson and loss of inner lobes on the lower lip.

The two groups at times have been considered congruent but there remains the
possibility that two or more ancestries occur, the Weckeliids from a Paraweck-
elia-Ceradocus ancestry and the Hadziids from a Melitid ancestry. As shown in
the Key to the Hadziids and Weckeliids, the groups are so contrived that elevation
to family level is impossible.

Key to Hadziids and Weckeliids

Meuich famus Of Uropod 3 with 2 articles .................+.: (Hadziids) 2
PeOuter ramus Of uropod 3 with | article ..........:....... (Weckeliids) 10

178

10.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Inner plate of maxilla 1 lacking medial setae, inner plate of maxilla 2

lacking oblique facial row of setae, medial margin setose or not ...... 3
Inner plates of maxillae 1-2 strongly setose medially, maxilla 2 with

oblique facial TOW/OlSetaAe* 4 TARE Bee a ee ne ae Se eee 4
Uropod 3 magniramous...2- 4.45... soos eee eee Paraweckelia (twice)
Uropods3iparviramousy Ae nei ae Se eee Psammoniphargus
Coxa’4 with largesposterodistalllobe) ‘A293. Geen ee ee Saliweckelia
Coxa 4cunlobedel). Pe SEPA RTA Sai Ce 5
Wrists of gnathopods 1—2 unlobed, their setae marginal .............. 6
Wrists of gnathopods 1-2 lobed, some of their lateral setae facial ..... 9
Telson shortened (uropod 3 variramous, gnathopods and telsonic spi-

nation like Metahadzia couplet below) .................. Metaniphargus
Nelsonof ordinary. lenethior elongates, 2 eae ee eee 7

Gnathopod | lacking medial setal brush on article 5, palm of male gnath-
opod 2 densely spinose, palm of female gnathopod 2 distinct, weakly
spinose, telson with medial spines, uropod 3 magniramous ... Metahadzia
Gnathopod | with medial setal brush, palm of male gnathopod 2 poorly
spinose, palm of female gnathopod 2 indistinct, telson lacking medial
spines, uropod)3: parviramous! 22222 is 2 eee VA, eee 8

. Palm of male gnathopod 2 densely setose, telson naked laterally ......

soos ew Siaw. Ghee OS ee eR eg Dulzura
Palm of male gnathopod 2 not densely setose, telson with lateral spines

BERS) ey ATED), MPL LT RET 9S bs ee ata ra oa Protohadzia
Telson with lateral spination, article 5 of gnathopod | elongate
6a Bat. de Soe TR ce Rae he ONS en ae Liagoceradocus
Telson naked laterally, article 5 of gnathopod 1 as long as6...... Hadzia
[In contradistinction to couplet 1, uropod 3 actually with vestigial article

2 on outer ramus], otherwise gnathopod 2 of both sexes of typical Ce-
radocid or Melitid form, hand well inflated, palm long, minutely serrate,
lacking Hadziid setae, wrist very short, strongly lobed, maxilla 2 lacking
oblique facial row of setae, inner plate of maxilla 1 with setae mostly
apical sis het: cals Loe eee Re ae ey ae ee Paraweckelia (twice)
[Like couplet 1, uropod 3 lacking article 2 on outer ramus], otherwise
gnathopod 2 not typical of Melitids or Ceradocids, hand either poorly
inflated or palm poorly defined in either sex or short, or not serrate, or
bearing Hadziid setae, wrist in either sex relatively elongate, maxillae
well setose medially, inner plate of maxilla 2 with oblique row of facial

Setar shied: Sach ulated ihe Mtge cee eg ce yak er ctik Salk Selene eee geen me fil
,'Coxay4 with posterodistal lobe: ga23 f2:Se. ee os Re oe eee 12
Coxa 4 without posterodistal lobe .....)\.05 454...) oe ee ee 13
. Inner lobes of lower lip present, palp of mandible l-articulate ... Weckelia
Inner lobes of lower lip absent, palp of mandible 3-articulate .........
PREG Pg ee RIMES EO EE OO Alloweckelia
. Coxa | not enlarged, right lacinia mobilis present ................... 14
Coxa | much larger than coxa 2, right lacinia mobilis absent ......... 16
. Wrist:of gnathopodsdobates wih. Wilk. Sa ee ee Mexiweckelia

Wrist.of enathopods not lobater= alee. he ee ee eee 15

VOLUME 95, NUMBER 1 179

ise Gnathopod | not merochelate’.. 0.5/0 eso... ee. Mexiweckelia particeps
PGnahiopod= | merochelate. ‘Yo 2:22 ie eta ee els Mayaweckelia
16. Mouthparts projecting anteriorly as far as article 2 of antennal ...... 17

— Mouthparts (except maxilliped) projecting less than halfway along article
| Git SIMUSTUOAL ST piel ORR Oe ee RAS ee ae Gare ne Ran ae Lt os) ae ern eee 18

17. Dactyl of maxilliped short, article 3 stout, inner plates of maxillae 1-2

elongate, outer plate of maxilla 1 with 7 spines, inner plate of maxilliped
CIPCIINY so ged! Saar io a ARR ear a ae ee a ae Holsingerius

— Dactyl of maxilliped elongate, article 3 thin, inner plates of maxillae 1-2

ordinary, outer plate of maxilla 1 with 14 spines, inner plate of maxilliped
aeRO NIA GLC Cl arg re ek a ettesoiie cond wee ones Be Golore el miagens here Texiweckeliopsis

18. Hand of gnathopod 1 like that of gnathopod 2, longer than wrist, gnatho-

pods of sexes alike, article 2 of pereopods 3-4 strongly expanded .....
i. Oe ins ny ate tepaitanaibe aun. 4 ala ciatey a yerd mua se Allotexiweckelia

— Hand of gnathopod | different from gnathopod 2, shorter than wrist,

gnathopods of sexes strongly distinct, article 2 of pereopods 3-4 thin ..
I Fok ee atl h eb oho art uot Neue tusudeteuksieroie, Ae Texiweckelia

Texiweckeliopsis, new genus

Type-species.—Texiweckelia insolita Holsinger, 1980.

Body slender, urosomites 1-3 each with 2, 4, and 2 dorsal spines respectively.
Lateral cephalic lobes truncate, without sinus. Eyes absent.

Antennae elongate, antenna | longer than 2, ratio of peduncular articles =
23:9:6, ventral margin of article | lacking spines. Accessory flagellum vestigial
or absent (smaller than in other Hadziids).

Mandibles and maxillae projecting forward as far as article 2 of antenna 1.
Mandibular right lacinia mobilis absent, palp absent. Labium without inner lobes,
not gaping. Maxillae medially setose, inner plate of maxilla 1 triangular, fully
setose medially, outer plate with 14-15 serrate spines, palps [?asymmetrical].
Plates of maxilla 2 slightly broadened, short, inner with oblique facial row of
setae. Inner plate of maxilliped extremely broad, outer plate medially setose;
article 3 of palp thin, with very small lobe forming weak apical chela, dactyl very
long, thin, with nail (maxilliped palp thus weakly prehensile).

Coxae of medium size to short, but coxa I larger than coxa 2, coxa 4 unlobed,
coxa 5 [?shorter than coxa 4]. Gnathopods subchelate, scarcely dimorphic sex-
ually, feeble; gnathopods 1-2 almost alike, mittenform, wrists elongate, broadly
lobate, setation facial, hands slightly shorter than wrists, narrow but not perfectly
linear or rectangular, slightly expanded in middle and then tapering towards palm,
palms weakly oblique, short, spines tiny or sparse, weakly bifid, Hadziid setae
few, elongate only on gnathopod 2 and sparse on palm, male gnathopods slightly
stouter than in female, especially on hands.

Pereopod 5 slightly shorter than 6—7, article 2 of pereopods 5-7 slightly ex-
panded, weakly lobate, dactyls with several setules on inferior margin, article 6
weakly setose or spinose.

Rami of uropods 1-2 subequally extended, uropod | with 2 basofacial spines,
comb of uropod 2 absent, outer ramus of uropod 2 dorsally naked. Uropod 3

180 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

elongate, magniramous, almost aequiramous, outer ramus |-articulate, apices of
rami weakly truncate, with several spines of which one elongate. Telson of or-
dinary length, cleft about two thirds, each lobe with 2 apical spines, main setules
M. 90.
Coxal gills 2-6, ovate, with long stalks, 2-articulate. Oostegites narrow.
Relationship.—Differing from Texiweckelia in the strong forward projection of
the mandibles and maxillae, and mittenform gnathopods. See Holsingerius.
Species.—insolita (Holsinger, 1980), Texas, hypogean, | species.
Etymology.—Name composed of Texiweckelia and ‘‘opsis’’ for “‘likeness.”’
Feminine.

Holsingerius, new genus

Type-species.—Texiweckelia samacos Holsinger, 1980.

Body slender, urosomites 1-3 each with 2, 2 and 4 dorsal spines respectively.
Rostrum obsolescent, lateral cephalic lobes sharply mammilliform. Eyes absent.

Antennae elongate, antenna | longer than 2, ratio of peduncular articles =
30:16:9, ventral margin of article 1 lacking spines. Accessory flagellum vestigial
or absent (smaller than in other Hadziids).

Mandibles and maxillae projecting forward as far as article 2 of antenna 1.
Mandibular right lacinia mobilis absent, palp absent. Labium without inner lobes,
gaping. Maxillae medially setose, inner plate of maxilla 1 enlarged, triangular,
fully setose medially, outer plate with 7 serrate spines, palps [?asymmetrical].
Inner and outer plates of maxilla 2 greatly elongate, inner with oblique facial row
of setae. Outer plate of maxilliped medially setose; article 3 of palp stout, apically
expanded and weakly chelate, dactyl of ordinary length, thin, with nail (palp
scarcely prehensile).

Coxae of medium size to short, but coxa I larger than coxa 2, coxa 4 unlobed,
coxa 5 [?shorter than coxa 4]. Gnathopods subchelate, sexually dimorphic, those
of female feeble; wrist of both pairs in both sexes of medium length, strongly
lobate, with facial setation; hand of female gnathopod | shorter than wrist, rect-
angular, palm short, weakly oblique, minutely spinose, male hand enlarged, palm
long, strongly oblique, well spinose; hand of female gnathopod 2 as long as wrist
(thus elongate), thin, rectangular, palm short, weakly oblique, minutely spinose
and with several Hadziid setae; anterior margin of hand lined with setae; hand
of male gnathopod 2 enlarged, palm long, rounded-oblique, densely spinose
(spines weakly bifid), with 2 Hadziid setae, dactyl very long. Article 2 of per-
eopods 3-4 scarcely expanded, poorly spinose. Pereopod 5 shorter than 6-7;
article 2 of pereopods S5—7 expanded, lobate, dactyls short, with several setules
on inferior margin, article 6 weakly setose or spinose.

Rami of uropods 2-3 subequally extended, uropod 1 with several basofacial
spines, comb of uropod 2 absent. Uropod 3 elongate, magniramous, almost ae-
quiramous, outer ramus l-articulate, apices of rami weakly truncate, with several
spines, of which one elongate. Telson scarcely elongate, cleft almost three-
fourths, each lobe with 3 apical spines and occasional lateral spine, main setules
MEGS:

Coxal gills 2-6, at least some ovate, with long stalks, 2-articulate. Oostegites
narrow [but full adult unknown].

VOLUME 95, NUMBER 1 181

Relationship.—Differing from Texiweckeliopsis in the ordinary dactyl of the
maxilliped, ordinary outer plates of maxillae and maxillipeds but the elongate
inner plates of the maxillae.

Species.—samacos (Holsinger, i980), Texas, San Marcos Well, hypogean, 1
species.

Etymology.—Named for the eminent authority on freshwater amphipods, Dr.
John R. Holsinger.

Paracalliopiidae, new family

Head, eyes, mouthparts, coxae 1-3, pereopods 3-6, uropods 1-2 ordinary.
Accessory flagellum vestigial. Coxa 4 poorly excavate posteriorly. Female
gnathopods feeble, mittenform, male gnathopods larger, gnathopod 2 enlarged,
wrist small, hand large (usually rotated inward on death), palm oblique. Pereopod
7 elongate, dactyl elongate and setose. Uropods 1-3 extending equally, peduncle
of uropod 3 slightly elongate, rami short, equal, lanceolate, outer l-articulate.
Telson laminar, entire. Urosomites 2-3 coalesced.

See Eusiridae, Pontogeneiidae, Calliopiidae, Oedicerotidae.

Male antennae with calceoli; antenna 2 longer than antenna |. Setosity of inner
plates on maxillae variable. Wrists of female gnathopods elongate, weakly to
strongly lobate posteriorly, lobes rounded and pointing terminad, hands slender,
palms oblique, short; wrists of male gnathopods not elongate, lobes pointing
almost perpendicular to axis of appendage, hands of ontogenetic appearance,
palmar defining corners softly rounded, palms of both gnathopods with long setae,
palm of gnathopod 2 also with stout spines; gnathopods of both sexes with patches
of pubescence. Pleon well developed, epimera large. Body not carinate.

Distinguished from Eusiridae, Pontogeneiidae, Calliopiidae, and Oedicerotidae
by the fused urosomites. Standing as an evolutionary grade between Calliopiidae
and Oedicerotidae by mixing the head, body form, and unspecialized mouthparts
of Calliopiidae with the elongate pereopod 7, elongate dactyl of pereopod 7,
setose dactyl of pereopod 7, slightly elongate peduncle of uropod 3, and rudiments
of fossorial adaptations in the slightly increased setosity of pereopods character-
istic of Oedicerotidae. The fused urosomites, however, show that the group is
divergent from the main evolutionary track.

Key to the Genera of Paracalliopiidae

1. Inner plates of maxillae densely setose medially ............. Paracalliope
— Inner plates of maxillae not setose medially ................. Indocalliope

Paracalliope Stebbing
Paracalliope Stebbing, 1899:210; 1906:279.—J. L. Barnard, 1972b:70.

Type-species.—Calliope fluviatilis Thomson, 1879 (original designation).

Inner plates of maxillae 1-2 densely setose medially. Uropods 1-2 normally
spinose.

Species.—australis (Haswell, 1880); fluviatilis Thomson (J. L. Barnard, 1972b);
karitane J. L. Barnard, 1972b; novaecaledoniae Ruffo and Vesentini-Paiotta, 1972;

182 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

novizealandiae (Dana) (?J. L. Barnard, 1972b); Dubious: fluviatilis of Chilton,
1921; New Zealand, Australia, middle IndoPacific to India; fresh and brackish
waters, marine littoral.

Nomenclatural changes.—Paracalliope fernandoi Wignarajah (1958) belongs
in the family Talitridae.

Indocalliope, new genus

Type-species.—Paracalliope indica K. H. Barnard, 1935.

Inner plates of maxillae not setose medially. Spines of uropods 1-2 sparse or
absent.

Species.—indica K. H. Barnard, 1935 (Nayar, 1959); India, brackish waters,
1 species.

Etymology.—Name composed from “‘India’’ and ‘‘Calliope.’’ Feminine.

Phoxocephalidae
Feriharpinia, new genus

Type-species.—Harpinia ferenteria Gurjanova, 1977.

Rostrum unconstricted. Eyes absent. Article 2 of antenna | short, ventral setae
widely spread. Article 1 of antenna 2 [?ensiform], article 3 with 3 facial setules,
facial spines on article 4 in [?1 row, ?all spines thin], article 5 ordinary. Right
mandibular incisor with [?3 teeth, right lacinia mobilis ?bifid, molar not triturative,
with ?3+ basally fused splayed spines; ?special spines; ?palpar hump small], apex
of palp article 3 oblique. Inner plate of maxilla | with 1 seta, palp biarticulate.
Maxillipeds ordinary, apex of palp article 3 not strongly protuberant, dactyl elon-
gate, apical nail [?distinct].

Gnathopods ordinary, small, [?dissimilar, gnathopod 2 ?weakly enlarged], ar-
ticle 5 of gnathopods 1-2 very short, free, palms oblique, hands ordinary, ova-
torectangular, poorly setose anteriorly.

Article 5 of pereopods 3-4 [? without posteroproximal setae, article 6 with ?thin
armaments]. Article 2 of pereopod 5 of [?narrow] form, articles 4—5 of pereopods
5—6 narrow; pereopod 7 ordinary, article 3 slightly enlarged, dactyl ordinary.

Epimera 1-2 [?without long facial brushes or posterior setae], epimeron 3 of
ordinary classification, bearing 4 or more long setae. Urosomite 3 [?without dorsal
hook].

Peduncle of uropod 1 [?without interramal spike, ?without major displaced
Spine, rami of uropods 1-2 ?not continuously spinose to apex, inner ramus of
uropod | with ?1 row of marginal spines]. Inner ramus of uropod 2 [?ordinary].
Uropod 3 [?ordinary], one of rami [?longer than peduncle], bearing [?article 2 on
outer ramus with ?2 apical setae]. Telson ordinary, but slightly elongate.

Relationship.—Like Harpinia but male armaments distinctive: instead of
brushes being present on article | of antenna | and article 1 of flagellum of antenna
1 and on articles 3-4 of antenna 2, and instead of article | of primary flagellum
on antenna | being enlarged and dominant, the male of Feriharpinia has a brush
of aesthetascs on article 3 of the peduncle of antenna 1, article 1 of the primary
flagellum is not grossly enlarged, and article 5 of antenna 2 has a row of large
dorsal calceoli, not found in Harpinia.

VOLUME 95, NUMBER | 183

Species.—ferenteria Gurjanova, 1977; Okhotsk Sea, west Kamchatka, 196-230
m, | species.

Etymology.—Name composed from “‘ferenteria’’ and related genus ‘‘Harpinia.”’
Feminine.

Proharpinia Schellenberg, (new composition)

Proharpinia Schellenberg, 1931:80 (Proharpinia antipoda Schellenberg, 1931,
monotypy).—J. L. Barnard, 1960:311.—Barnard and Drummond, 1978:532.

Rostrum unconstricted. Eyes present.

Article 2 of antenna | short, ventral setae almost confined apically (by fiat).
Article 1 of antenna 2 not ensiform, article 3 with 4—5 facial setules, facial spines
on article 4 in 1 main row, all spines thin, article 5 short. Right mandibular incisor
with [?3 teeth, right lacinia mobilis ?bifid, ?weakly flabellate, ?molar not tritu-
rative, with ?3 splayed spines]; palpar hump small, apex of palp article 3 oblique.
Inner plate of maxilla 1 naked, palp biarticulate. Maxillipeds ordinary, apex of
palp article 3 not strongly protuberant, dactyl body not elongate, but apical nail
distinct and elongate.

Gnathopods small, scarcely dissimilar, gnathopod 2 weakly to moderately en-
larged, article 5 of gnathopods 1-2 very short, free on gnathopod 1, cryptic on
gnathopod 2, palms oblique, hands of gnathopods 1-2 ovatorectangular, elongate,
poorly setose anteriorly.

Article 5 of pereopods 3—4 with posteroproximal setae, article 6 with thin
armaments. Article 2 of pereopod 5 of narrow form, articles 4-5 of pereopods
5—6 medium to narrow; pereopod 7 ordinary, but article 3 enlarged, dactyl ordi-
nary.

Epimera 1-2 without long facial brushes or posterior setae, epimeron 3 of
ordinary classification, bearing 3 or more long setae. Urosomite 3 without dorsal
hook.

Peduncle of uropod | without interramal spike, without major displaced spine,
rami of uropods 1-2 continuously spinose to apex, with subapical spines or nails,
inner ramus of uropod | with | row of marginal spines. Inner ramus of uropod
2 ordinary. Uropod 3 ordinary, | of rami longer than peduncle, bearing article 2
on outer ramus, with vestigial or no apical setae. Telson ordinary.

Variants.—Proharpinia stephenseni, the second species of the genus, is re-
tained here only provisionally; it differs from the type in the presence of setae on
the inner plate of the maxilliped and the absence of apical spination on the rami
of uropods 1-2. However it matches the type in the telson, and loss of strong
setation on article 2 on the outer ramus of uropod 3.

Relationship.—Differing from Heterophoxus in the absence of an ensiform pro-
cess on antenna 2; more plesiomorphic than Torridoharpinia in spination of uro-
pods 1-2 and normal condition of outer ramus of uropod | (not shortened), and
normal armament of telson but more apomorphic in loss of setae on inner plate
of maxilla 1. Proharpinia hurleyi and P. tropicana are removed to Torridohar-
pinia below.

Species.—See J. L. Barnard, 1960; antipoda Schellenberg, 1931; stephenseni
Schellenberg, 1931; Magellan area, Falkland Islands, 2-274 m.

184 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Torridoharpinia, new genus

Type species.—Proharpinia hurleyi J. L. Barnard, 1958, 1960.

Rostrum unconstricted. Eyes present.

Article 2 of antenna | short, ventral setae ventrally spread or almost confined
apically. Article | of antenna 2 not ensiform, article 3 with 4 facial setules, facial
spines on article 4 in | main row, all spines thin, article 5 short. Right mandibular
incisor with 3 teeth, right lacinia mobilis bifid, flabellate, molar not triturative,
with 2 splayed spines; palpar hump small, apex of palp article 3 oblique. Inner
plate of maxilla 1 with 2-4 (type) setae, palp biarticulate. Maxillipeds ordinary,
apex of palp article 3 not strongly protuberant, dactyl elongate, apical nail dis-
tinct.

Gnathopods small, dissimilar, gnathopod 2 weakly to moderately enlarged,
article 5 of gnathopods 1-2 short, free on gnathopod 1, cryptic on gnathopod 2,
palms oblique, hands ovatorectangular, broadened on gnathopod 2, poorly setose
anteriorly.

Article 5 of pereopods 3-4 with posteroproximal setae, article 6 with thin ar-
maments. Article 2 of pereopod 5 of narrow form, articles 4-5 of pereopods 5—6
medium to narrow; pereopod 7 ordinary, article 3 enlarged, dactyl ordinary.

Epimera 1-2 without long facial brushes or posterior setae, epimeron 3 of
ordinary classification, bearing 3 or more long setae. Urosomite 3 without dorsal
hook.

Peduncle of uropod | without interramal spike, without major displaced spine,
rami of uropods 1|—2 not continuously spinose to apex, inner ramus of uropod 1
only in male with 2 rows of marginal spines. Inner ramus of uropod 2 ordinary.
Uropod 3 ordinary, one of rami longer than peduncle, bearing article 2 on outer
ramus, with | or 2 (type) apical setae. Telson ordinary, but 1 apical element stout
(contrast Proharpinia).

Relationship.—Differing from Proharpinia in the lack of ramal spines on female
uropod 2, the shortened outer ramus of uropod 1, the presence of 1—2 long apical
setae on article 2 of the outer ramus on uropod 3, the presence of a stout spine
on each lobe of the telson and the presence of setae on the inner plate of maxilla
1. Proharpinia has only thin setae or setules on the telson.

Species.—hurleyi J. L. Barnard, 1958, 1960 (=stephenseni of Hurley, 1954, not
Schellenberg, 1931); tropicana J. L. Barnard, 1960; New Zealand, Auckland Is-
lands, Campbell Island, Galapagos Islands, 0-46 m, 2 species.

Etymology.—Name composed from “‘torrid’’ and ‘‘Harpinia.’’ Feminine.

Pseudamphilochidae Schellenberg, 1931

Only coxa 4 slightly broadened, coxae 2-4 with continuous margins overlap-
ping, not rabbeted, coxae 1-2 not hidden, coxa | not reduced, no anterior coxa
hidden. Telson cleft.

Upper lip weakly excavate but unnotched. Mandibular molar vestigial or ab-
sent. Lower lip with well developed inner lobes.

Palp of maxilliped lacking process on article 3.

Outer ramus of uropod | shortened. Peduncle of uropod 3 not greatly elongate,
rami almost twice as long as peduncle.

VOLUME 95, NUMBER 1 185

Remarks.—This family has not been properly heralded in the literature before
and is brought to light from a provisional proposal (Schellenberg, 1931: 92).

Pseudamphilochus Schellenberg
Pseudamphilochus Schellenberg, 1931:92.

Type-species.—Pseudamphilochus shoemakeri Schellenberg, 1931 (monotypy).

With familial characters.

Gnathopods of medium size, wrists short, weakly lobate, not carpochelate,
hands broad, scarcely 1.4 times as long as broad, palms subtransverse.

Species.—shoemakeri Schellenberg, 1931; South Georgia, littoral.

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Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar,
Supplementary Report 17:229-300, 8 pls.

Wignarajah, S. 1958. Paracalliope fernandoi sp. nov., a new fresh-water amphipod from Ceylon.—
Ceylon Journal of Science, Biological Science, 1:115—116, 2 pls.

PROC. BIOL. SOC. WASH.
95(1), 1982, pp. 188-193

HEMIODOPSIS OCELLATA, A NEW HEMIODONTID
CHARACOID FISH (PISCES: CHARACOIDEA)
FROM WESTERN SURINAM

Richard P. Vari

Abstract.—Hemiodopsis ocellata, an open water species occuring in the biack
water drainages of western Surinam, is described from several localities within
the Corantijn River system basin. The pattern of zig-zag lines on the dorsal por-
tion of the body appears to be unique in the genus.

Introduction

The new species described herein, Hemiodopsis ocellata, is widespread in the
black waters of the Corantijn River system of western Surinam. Habitats of in-
dividuals captured or observed in the wild show that during low water levels this
Species 1s primarily an inhabitant of the main river channels and larger open pools
and streams rather than the shaded creeks and smaller streams of the rain forest.
During high water levels, in contrast, H. ocellata apparently travels up smaller
tributaries into the flooded rain forest to feed and reproduce. The species has
been taken most often in the more readily sampled still or slowly flowing waters.
Nonetheless, the capture of a single specimen in the rapidly-flowing waters of
Dalbana Creek and observations over several seasons indicate that H. ocellata
also occurs in waters with high flow velocities. This species is usually found in
association with the much more abundant Hemiodopsis goeldii, which it closely
resembles in overall body form, maximum adult size, and pigmentation pattern
(see Bohlke, 1955, fig. 7).

Hemiodopsis ocellata, new species
Fig. 1, Table 1

Holotype.—National Museum of Natural History (USNM) 221175, 170.0 mm
standard length (SL), collected by R. P. Vari, 7 December 1979 in the main stream
of Dalbana Creek, approximately 150 m upstream of its junction with the Ka-
balebo River, Nickerie District, Surinam (approx. 4°47'N, 57°29’'W).

Paratypes.—1 specimen, USNM 225592, 141.5 mm SL, collected by R. P. Vari
and L. R. Parenti, 8 September 1980, in a slow-flowing side channel of the Cor-
antijn River about 180 km from its mouth, Nickerie District, Surinam (approx.
5°08’N, 57°18’W); 2 specimens, USNM 225593, 104.7-113.0 mm SL [1 cleared
and counterstained for cartilage and bone] collected by R. P. Vari and L. R.
Parenti, 17 September 1980, in a still pool near ‘‘Camp Hydro’’ on an island in
the middle of the Corantijn River, Nickerie District, Surinam (approx. 4°22'N,
57°58'W); 4 specimens, USNM 225594, 129.0-142.0 mm SL, collected by H. M.
Madarie, 15 May 1980, in a small creek and the surrounding flooded rain forest
of the Corantijn River, Nickerie District, Surinam (approx. 5°32'N, 57°10’W).

Diagnosis.—Within the family Hemiodontidae the new species is assignable to

VOLUME 95, NUMBER 1 189

Fig. 1. Hemiodopsis ocellata, new species, holotype, USNM 221175, 170.0 mm SL.

Hemiodopsis based on the very slightly protractile jaws (in contrast to the highly
protractile jaws of Bivibranchia and Argonectes), multicuspidate dentition limited
to the upper jaw (in contrast to unicuspidate teeth in both jaws in Michromis-
chodus), a dorsal fin of moderate height (in contrast to an elongate, anteriorly
filamentous dorsal fin in Pterhemiodus), and a moderate vertical gradation in
scale size (in contrast to a pronounced vertical gradation in scale size in He-
miodus). Within Hemiodopsis (see Géry 1963 for a discussion of the redefinition
of the genus) the 88—96 lateral line scales occurring in H. ocellata readily distin-
guish it from the majority of the species in the genus (H. gracilis, H. goeldii, H.
fowleri, H. thayeri, H. semitaeniata, H. ternetzi, H. immaculata, H. rodolphoi,
and H. parnaguae) which as a unit demonstrate a range from 42 to 83 lateral line
scales, and from H. microlepis and H. argentea which have a higher number of

Table 1.—Morphometrics of Hemiodopsis ocellata, new species. Standard length is expressed in
mm; measurements | to 11 are percentages of standard length; 12 to 15 percentages of head length.

Paratypes (7)

Holotype Range Average

Standard length 170.0 113.0-141.5 129.6

1. Greatest body depth 29% 27.4—29.8 291

2. Snout to dorsal-fin origin 49.2 42.8-54.0 49.3

3. Snout to anal-fin origin 80.9 80.7-82.9 81.6

4. Snout to pelvic-fin origin 52.8 52.9-54.0 53.4
5. Snout to anus 78.0 77.0-77.9 78.1

6. Origin of rayed dorsal to hypural joint 56.0 53.9-56.3 55.0

7. Least depth of caudal peduncle 10.1 9.1-11.2 10.0

8. Pectoral-fin length 20.0 19.0-21.2 19.7

9. Pelvic-fin length 21.8 23-241 22.6

10. Dorsal-fin height 26.4 25.1—30.2 Dey
11. Head length 26.1 25.0-26.4 25.6
12. Orbital diameter 30.0 29.0-32.4 30.1
13. Snout length 31.0 29.2—34.2 31.6
14. Postorbital length 38.4 37.3-40.0 38.8
15. Interorbital width 36.8 34.9-38.1 36.1

190 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

585 a7”

58° 57k

Fig. 2. Map of the lower Corantijn River basin region, Surinam and Guyana, showing collecting
localities for the following specimens of Hemiodopsis ocellata (dotted line depicts the road to Camp
Matapi and Camp Amotopo): 1, collection locality of holotype, USNM 221175; 2, collection locality
of 2 paratypes, USNM 225593; 3, collection locality of 1 paratype, USNM 225592; 4, collection
locality of 4 paratypes, USNM 225594.

VOLUME 95, NUMBER 1 191

pored lateral line scales (110 to 112 and 120 to 125 respectively). The only He-
miodopsis species having a lateral line scale count approximating that of H.
ocellata is H. parnaguae of the upper Parnaiba River system (80-83 lateral line
scales; Eigenmann and Henn 1916:87). However, in addition to the differences
in the number of pored lateral line scales, these species are distinguishable by
the hyaline anal fin of H. ocellata which contrasts with the presence of a distal
black band on the fin in H. parnaguae (see Eigenmann and Henn 1916, Pl. 17).
Furthermore, H. parnaguae lacks the distinctive dorsal zig-zag patterning char-
acteristic of H. ocellata.

Description.—Table 1 gives morphometrics of the holotype and paratypes.
Body relatively slender, slightly compressed laterally. Greatest body depth in
region of origin of rayed dorsal fin. Dorsal profile of body gently curved from tip
of snout to interorbital region, slightly convex or straight from that area to origin
of rayed dorsal fin. Body profile at base of rayed dorsal fin straight, posteroven-
trally sloped. Dorsal profile of body nearly straight from rear of insertion of rayed
dorsal fin to upper margin of caudal peduncle. Ventral profile of head smoothly
convex from tip of lower lip to below base of pectoral fin; nearly straight from
that point to origin of anal fin.

Head relatively small, snout obtuse with mouth subinferior. Anterior margin
of lower jaw cresent-shaped. Upper jaw moderate, very slightly protractile, pos-
terior margin of maxilla extends slightly posterior of a vertical through posterior
border of rear nostril. Nostrils approximate, anterior opening round, posterior
cresent-shaped. Eye relatively large. An extensive, horizontally ovoid ‘‘adipose
eyelid’ (a thick transparent connective tissue layer) extends from posterior mar-
gin of rear nostril to middle of opercle (less developed posteriorly in smaller
specimens). Adipose eyelid with an ovoid, vertically-elongate opening overlying
pupil. Fronto-parietal fontanel extensive, extending into rear of ethmoid. Parietals
completely separated, frontals in contact only at epiphyseal bar. 4 branchiostegal
rays on each side; | on posterior ceratohyal, 3 on anterior ceratohyal. Osteolog-
ical characters are overall very similar to those of Hemiodus (=Hemiodopsis)
semitaeniata as illustrated by Roberts (1974). Vertebrae 39 or 40 including those
of Weberian apparatus and counting fused PU,+U, as a single element.

Lower jaw edentulous, anterior edge sectorial. Functional teeth in a single
series in upper jaw. All teeth multicuspidate; number of cusps on each tooth
decreasing from 11 on medial teeth on 7 on lateral teeth. Sixteen to 18 teeth along
each side of upper jaw, 10 maxillary and 8 premaxillary teeth on each side of jaw
in cleared and stained specimen. Teeth extending along entire anterior edge of
maxilla (see Roberts, 1974, fig. 9 for comparable situation in Hemiodopsis sem-
itaeniata). One or two rows of partially formed replacement teeth internal to
functional tooth row. Premaxillary replacement tooth rows embedded in flesh on
inner surface of premaxilla; maxillary replacement teeth arranged in a shallow
maxillary replacement-tooth trench. Dermopalatine and ectopterygoid edentu-
lous. Ceratobranchial 5 bearing a relatively narrow band of posteriorly-directed
teeth along medial and posteromedial borders. Upper pharyngeal tooth-plates 4
and 5 with bands of small, conic teeth. Ceratobranchials 2, 3, and 4 and hypo-
branchial 3 bearing ventrally-directed processes extending lateral to ventral aorta.
Gill-rakers expanded distally, fan-shaped, with a series of pointed, digitiform
processes along distal edges. Number and size of distal gill-raker processes in-

192 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

creasing towards ceratobranchial-epibranchial joint. Gill-rakers on first gill arch
38 on epibranchial and 50 on ceratobranchial in cleared and counterstained spec-
imen.

Scales cycloid, thin. Pored lateral line scales between supracleithrum and hy-
pural joint 93 in holotype (88 in | paratype, 92 in 1 paratype, 93 in 2 paratypes,
95 in 2 paratypes, 96 in | paratype). Five to 7 pored lateral line scales extending
beyond hypural joint onto caudal fin. Scales above lateral line in a transverse
series to origin of rayed dorsal fin 22 in holotype (22 in 3 paratypes, questionably
21 in 4 partially descaled paratypes). Scales below lateral line in a transverse
series to origin of anal fin 14 in holotype (12 in | paratype, 13 in 1 paratype, 14
in 2 paratypes, 15 in 1 paratype, 2 paratypes descaled in region). Body squamation
extending onto base of caudal rays. Low sheath of scales along base of rayed
dorsal fin and anterior portion of anal fin. Axillary process of pelvic fin composed
of a series of scales. Size of body scales only slightly graduated in a vertical
series.

Rayed dorsal fin pointed but not filamentous anteriorly; second unbranched
and first branched rays longest, subequal. Dorsal-fin rays 11,9 in all specimens.
Adipose dorsal fin moderate, unscaled; length about two-thirds diameter of orbit.
Anal fin falcate, anterior branched rays more than twice as long as posteriormost
branched rays. Anal-fin rays 11-9 in holotype (1i,9-i in | paratype, 1-9 in 2 para-
types, 11-10 in 4 paratypes), first 2 unbranched anal rays very short. Pectoral fin
pointed, reaching two-thirds distance to a vertical through insertion of pelvic fin.
Pectoral-fin rays 1-16-11 in holotype (i-16-ii in 3 paratypes, i-16-111 in 2 paratypes,
i-17-i1 in 2 paratypes). Pelvic fin pointed, reaching three-quarters distance to anus.
Pelvic-fin rays i-10 in holotype (i-10 in 6 paratypes, 1-11 in | paratype).

Coloration in alcohol.—Overall coloration silvery. Head dark dorsally, a much
more intensely pigmented spot about size of pupil located on anterodorsal corner
of opercle and surrounding region. Fleshy lip of premaxilla and maxilla dark.
Body darker dorsal to lateral line, silvery-white ventrally. A diffuse dark lateral
band slopes slightly posteroventrally from dorsal portion of opercle to base of
caudal fin. Body band interrupted by an intensely pigmented, longitudinally elon-
gate, ocellated spot. Spot straddling but located largely dorsal to lateral line,
extending from slightly behind a vertical through rear of dorsal-fin insertion nearly
to a vertical through posterior limit of pelvic fin; spot extending about 12 scales
longitudinally and 7 vertically. Unpigmented region surrounding spot approxi-
mately 5S or 6 scales wide. Dorsal portion of body above lateral line marked by
31 to 35 dark, strongly-angled bars having the form of a “‘V”’ on its side with
angle of flexure directed posteriorly. Each bar about one and one-half or two
scales wide. Pigmentation pattern less apparent anteriorly and posteriorly. A
series of randomly arranged spots on sides of body below lateral line, each spot
about size of exposed portion of scales in that region. Rayed dorsal fin dusky,
with no apparent pattern. Adipose dorsal fin dusky. Anal fin clear. Pectoral fin
with narrow bands of light brown chromatophores outlining unbranched lateral
fin ray and first branched ray. Anal fin with series of pale brown chromatophores
outlining distal portions of rays on second through fourth branched anal rays.
Lower lobe of caudal fin with a dark stripe continuous with lateral band on body.
Procurrent rays and first and second principal rays of lower caudal lobes unpig-
mented. Procurrent rays of upper caudal lobe outlined by dark chromatophores.

VOLUME 95, NUMBER 1 193

Coloration in life. —Overall coloration silvery golden. Iris intense gold. Silvery
coloration somewhat masking lateral body band and dorsal bars. Fins pinkish or
light red.

Relationships.—Our present understanding of relationships within Hemiodop-
sis are quite poor. Indeed, no characters supporting an hypothesis of monophyly
of the genus have been proposed. Within Hemiodopsis as presently delimited, H.
ocellata is most similar to although necessarily most closely related to H. par-
naguae.

Etymology.—Ocellata, from the Latin for little eyes, in reference to the lateral
eye-like body spot.

Acknowledgments

The specimens that served as the basis for this paper were collected during
surveys carried out as a part of preimpoundment studies associated with the
Kabalebo Hydroelectric Project. The assistance of Drs. M. P. Panday-Verheuvel
and S. Niekoop is gratefully acknowledged as is field assistance by Dr. L. R.
Parenti, Mr. H. M. Madarie, Ms. S. Engel, and Mr. S. Silos. This manuscript
benefited from critiques by L. R. Parenti and S. H. Weitzman. Figure | is by
Susan L. Jewett.

Literature Cited

Bohlke, J. 1955. Studies on fishes of the family Characidae.—No. 10. Notes on the coloration of the
; species of Hemiodus, Pterhemiodus, and Anisitsia, with the description of a new Hemiodus

from the Rio Negro at the Brazil-Colombia border.—Notulae Naturae 278:1-15.

Eigenmann, C. H., and A. W. Henn. 1916. Description of three new species of characid fishes.—
Annals of the Carnegie Museum 10:87-90.

Gery, J. 1963. Sur la nomenclature et la systematique du genre Hemiodus Miiller (Pisces, Chara-
coidei).—Bulletin Museum National d’Histoire Naturelle, Series 2, 35:589-60S.

Roberts, T. R. 1974. Osteology and classification of the Neotropical characoid fishes of the families
Hemiodontidae (including Anodontinae) and Parodontidae.—Bulletin of the Museum of Com-
parative Zoology 146:411-472.

Department of Vertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
95(1), 1982, pp. 194-197

EVIDENCE OF ONTOGENETIC SETAL CHANGES
IN HETEROMASTUS FILIFORMIS
(POLYCHAETA: CAPITELLIDAE)

Thomas J. Fredette

Abstract.—Heteromastus filiformis undergoes ontogenetic setal replacement in
the fourth and fifth setiger. Setigers 4 and 5 initially bear hooded hooks which
are gradually lost and replaced by capillary setae.

The possible occurrence of ontogenetic setal changes in the polychaete Het-
eromastus filiformis was first observed in samples taken during a colonization
experiment in the lower York River, Virginia (Fredette 1980). The first sample
set (0.25 mm sieve), taken shortly after deployment of azoic substrates, contained
newly set juveniles of several different species of benthos, including a capitellid
polychaete. The general body morphology of these newly set capitellids was
similar to that of the 2 locally common species, Heteromastus filiformis and
Mediomastus ambiseta. However, the specific characters fit neither. The juve-
niles had capillary setae on the first 3 setigers while the remaining setigers bore
hooded hooks. This is quite unlike H. filiformis which has the first 5 anterior
setigers with capillary setae and M. ambiseta which has capillary setae in the
first 4 anterior setigers and the last several notopodia. Specific taxonomic place-
ment of these juvenile capitellids did not seem possible.

3 CAPILLARY SETIGERS

4 CAPILLARY SETIGERS

5S CAPILLARY SETIGERS

INDIVIDUALS
uw
oO

6)
O lO 15 34 73 144

TIME IN DAYS
Fig. 1. Progression of setal changes observed in a population of H. filiformis colonizing azoic
substrates in the York River, Virginia. Volume of area indicates percent of population exhibiting the
indicated character.

VOLUME 95, NUMBER 1 195

Fig. 2. SEM micrographs of juvenile capitellids: A, Anterior end of a juvenile capitellid with both
capillary and hooded hook setae on fifth setiger; B, Mixed setal bundle of a juvenile capitellid. Setal
bundle in background bears all capillary setae.

Successive samplings taken during the colonization process revealed serial
changes in setal pattern in the population of newly set capitellids (Fig. 1). The
individuals from these later samples constituted a continuum from the 3-capillary
setiger stage to a 5-setiger stage. The continuum was complete with intermediate
Stages (individuals with setal bundles containing both capillary setae and hooded
hooks) (Fig. 2) and a gradation of sizes. The graph shown in Figure | is based on
the assumption that all the juveniles are H. filiformis. However, individuals that
were obviously M. ambiseta also occurred in the samples from day 73 and day
144. The large time gaps in the sampling series made a final conclusion on the
specific placement of these juveniles difficult.

Additional documentation of this phenomenon was obtained by examining sam-
ples from an ongoing benthic study in the lower York River in which weekly
samples are being taken (Diaz, unpublished data). It was felt that these samples
would clarify the progression of setal changes and help to better isolate the ap-

196 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

100 2!| APRIL 28 MAY
n=2 n=60
% 50
100
28 APRIL 2 JUNE
n=O n=6!1
ou 50
ee 5 MAY 9 JUNE
n=7 n=94
% 50
100
13 MAY 16 JUNE
n=25 n= 132
% 50

100
19 MAY 23 JUNE
n=28 n=73
fy OO
AE Br ee Stier FP Ae Be Ceap ee F

Fig. 3. Weekly distribution of setal pattern in a field population of capitellids. A. 3 anterior
capillary setigers; B, 4th setiger with both capillary and hooded hook setae; C, 4 anterior capillary
setigers (no posterior capillaries); D, 5th setiger with both capillary and hooded hook setae; E, 5
anterior capillary setigers (H. filiformis pattern); F, 4 anterior capillary setigers, posterior capillaries
(M. ambiseta pattern). Areas indicated are percents of total population.

pearance of M. ambiseta and H. filiformis in the benthic community. Individuals
in these samples were placed into 6 classes: (A) 3 anterior capillary setigers, (B)
fourth setiger with mixed capillaries and hooded hooks (no posterior capillaries),
(C) 4 anterior capillary setigers (no posterior capillaries), (D) fifth setiger with
mixed capillaries and hooded hooks, (E) H. filiformis setal pattern, (F) M. am-
biseta setal pattern.

The occurrence of M. ambiseta in these samples was sporadic, in low densities
and seemed to bear no relationship to the development of the juvenile capitellids
(Fig. 3). Also, a few small individuals of M. ambiseta were found that were
similar in size to the other juvenile capitellids, indicating that M. ambiseta may
have the same setal pattern in juveniles and adults.

The other five classes appear in a pattern similar to the age-class distributions
one expects from a population with synchronous breeding and recruitment (Fig.
3). Recruitment occurs in early May and the population quickly passes through
the various setal changes until 7 weeks later when all the individuals exhibit the
typical H. filiformis pattern.

VOLUME 95, NUMBER 1 197

Table 1.—Width measurement (um) of fifth setiger, means or ranges.

Class Width
3 capillary setigers 116.2, a = 4
4th mixed 185-65) n— 9
4 capillary setigers QU. 1, 1 = 7
Sth mixed S12 — 6
H. filiformis (adults) 300-600
juvenile M. ambiseta 140-200

To illustrate the progression of sizes that occur, several individuals from each
class were measured by recording the width of the fifth setiger. The juvenile M.
ambiseta were also measured for comparison (Table 1). As indicated earlier these
individuals are similar in size to the 3-capillary setiger individuals. The juvenile
capitellids showed a steady progression of sizes and change to a 5-capillary setal
pattern.

That H. filiformis undergoes an ontogenetic development from a 3 anterior
capillary stage to a 5 anterior capillary stage is strongly indicated by the evidence
presented. Setigers 4 and 5 initially bear hooded hooks which are gradually lost
and replaced by capillary setae. Mediomastus ambiseta appears not to undergo
such a morphological change. Further substantiation of the observed setal
changes could be facilitated by examination of cultured populations of these two
capitellid species.

The separation of M. ambiseta and H. filiformis in benthic samples (0.5 mm
sieve) by anterior capillary setiger counts alone (see keys of Ewing and Dauer
1981, Fauchald 1977) is not sufficient. Examination for posterior notopodial cap-
illaries is necessary.

Acknowledgments

I would like to express my thanks to Dr. Robert J. Diaz for the loan of several
samples and critical review of this manuscript, Dr. Frank O. Perkins for advice
on SEM techniques and preparations, and Michael J. Kravitz and Stephanie A.
Vay for valuable suggestions and discussion.

Contribution No. 1039 from the Virginia Institute of Marine Science, The Col-
lege of William and Mary.

Literature Cited

Ewing, R. M., and D. M. Dauer. 1981. A new species of Amastigos (Polychaeta: Capitellidae) from
the Chesapeake Bay and Atlantic coast of the United States with notes on the Capitellidae of
the Chesapeake Bay.—Proceedings of the Biological Society of Washington 94: 163-168.

Fauchald, K. 1977. The polychaete worms, definitions and keys to the orders, families and genera.—
Natural History Museum Los Angeles County, Science Series 28:1-190.

Fredette, T. J. 1980. Macrobenthic colonization of muddy and sandy substrates in the York River,
Virginia. M.S. thesis, The College of William and Mary, Williamsburg, Virginia, 62 pp.

Department of Invertebrate Ecology, Virginia Institute of Marine Science and
School of Marine Science, The College of William and Mary, Gloucester Point,
Virginia 23062.

PROC. BIOL. SOC. WASH.
95(1), 1982, pp. 198-202

A NEW SPECIES OF OPHIOGOMPAUS (INSECTA:
ODONATA: GOMPHIDAE) FROM THE WESTERN
HIGHLAND RIM IN TENNESSEE

Jerry A. Louton

Abstract.—Ophiogomphus bouchardi, a new species of Gomphidae, is de-
scribed from the Western Highland Rim of central Tennessee. The description
includes both sexes and the exuviae of the holotype male. Affinities of the new
Species apparently lie with O. carolinus and O. mainensis.

While collecting crayfish on the Western Highland Rim in Dickson Co., Ten-
nessee, in 1971, Dr. Raymond W. Bouchard collected a single female nymph of
Ophiogomphus which he subsequently succeeded in rearing. The reared female
and exuviae were placed in the University of Tennessee collection of aquatic
insects (as O. mainensis) and there remained until examined by me in 1976. The
female and nymphal skin were at once seen to be distinct from other species of
the genus by several outstanding characters. I now have a series of seventeen
reared adults to support the original supposition of distinctness. Dr. Bouchard
has kindly consented to the inclusion of his 1971 collection as part of the type
series. It is with great pleasure that I name this new species in recognition of Dr.
Bouchard’s many contributions to aquatic biology.

Ophiogomphus bouchardi, new species
Figs. 1-13

Material.—Holotype male, TENNESSEE, Dickson Co., Will Hall Creek at
U.S. Hwy. 70 (entrance to Montgomery Bell State Park), nymph collected 21
Mar 1981, adult emerged 13 Apr 1981; allotype female, same location and date,
adult emerged 14 Apr 1981; paratypes, same location and date, 5 males and 3
females (all reared); same location, 1 female, nymph collected 26 Mar 1971, date
adult emerged unknown. Additional material not in type series, same location
and date, 3 reared males and 2 reared females; same location, 25 Sep 1980, 5 final
instar nymphs; Lewis Co., Litthe Swan Creek at Natchez Trace Parkway, 20 Mar
1979, | final instar nymph. All types will be placed in the Florida State Collection
of Arthropods, Gainesville, Florida.

Diagnosis.—Dorsal stripes of synthorax absent or vestigial; face not cross-
striped with black at sutures; yellow of tibiae restricted to one-fifth length of
tibiae; lateral process of male epiproct not strongly projected; anterior occipital
horns of females erect, separated by a distance equal to their height; posterior
occipital horns of females vestigial; prementum of nymphs narrowed anteriorly,
ligula narrow and strongly projected.

Description of holotype male.—Total length 47.4 mm, abdomen 34.1 mm, hind
wing 28.2 mm. Form and coloration generally typical for eastern North American
species.

VOLUME 95, NUMBER 1 199

= : Mee:

Pawel eae

4 lO l2 I3

Figs. 1-13. Ophiogomphus bouchardi, n. sp.: 1, Dorsum of synthorax of holotype male; 2, Lateral
view of synthorax of holotype male; 3-5, Extremes in form of occipital horns of females; 6, Anterior
hamule of holotype male; 7, Posterior hamule of holotype male; 8, Penis and vesicle of holotype
male; 9, Lateral view of terminal abdominal appendages of holotype male; 10, Dorsal view of terminal
abdominal appendages of holotype male; 11, Right antenna of exuviae of holotype male; 12, Ventral
view of subgenital plate of allotype female; 13, Prementum of exuviae of holotype male with inset of
detail of ligula.

200 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Anterior of frons yellow-green without cross stripes at sutures, dorsum nar-
rowly infuscated with brown between antennal bases, six black granules antero-
laterally, entire frons covered with black hairs. Vertex blackish across anterior
half, yellowish at hairy prominences, brownish on posterior half. Occiput yellow
with dense line of black hairs at crest, scattered black hairs on posterior surface.

Prothorax with two broad longitudinal black bands on dorsum. Synthorax light-
ly striped with brown; brown stripes adjacent to dorsal crest vestigial, only a
faint brown streak adjacent to middle prominence of crest (Fig. 1); second (an-
tehumeral) stripe of thorax present, continuous with brown of katepisternum
below, upper end not connected to brown of antealar ridge; stripe of second
plural suture (humeral) complete, wider in upper half and constricted somewhat
just as it joins antealar ridge; middle suture with faint stripe ending at spiracle;
stripe of third pleural suture absent except for faint streak on upper one-third
(Fig. 2). Femora greenish; streaked with black that is apically coalesced; tibia
and tarsi all black except for small narrow proximal spot of green on outer surface
of tibia.

Abdominal segments with dorsal yellow spots, wide on segments two and three,
narrowing to segment eight, wider on segments nine and ten; in lateral view
abdominal segments dark in upper one-half, gray-green below with black en-
croaching down into light areas along supplementary transverse carinae and as
dark postero-lateral spots; light areas of segments seven to ten tinged with yellow;
auricles yellow-green with brownish crescent behind. Male genitalia black-tipped,
vesicle shining black; anterior hamule bifid, tip claw-like (Fig. 6); posterior ham-
ule tapering, bent anteriorly forming smooth curve (Fig. 7); penis with moderate
flagella (Fig. 8). Terminal abdominal appendages yellow; superior appendages
taper to sharp tip, in dorsal view bowed in middle and proximate at bases and
tips (Fig. 10), in lateral view appendages arched in middle (Fig. 9), black denticles
scattered on ventral surface near tips; inferior appendage bifid to one-half its
length, lateral prominences are low tubercles placed near midlength of appendage.

Description of allotype female.—Total length 46.7 mm, abdomen 31.9 mm,
hind wing 30.2 mm. Similar in general form and coloration to holotype male
except: occiput with pair of large erect sharp-tipped horns separated by distance
equal to their length (Fig. 5) and surrounded by black hairs; no vestige of brown
stripes adjacent to dorsal crest of synthorax; posterolateral black spots on ab-
dominal segments continue anteriorward as streaks on middle abdominal seg-
ments; vulvar lamina as long as ninth abdominal sternite (Fig. 12).

Exuviae of holotype male.—Total length 26.3 mm. Exuviae light brown with
coarse pigmented cuticular granules.

Antennae broad and flat, two and one-fourth times longer than wide (Fig. 11).
Prementum narrowed anteriorly; palpal lobes short, two-fifths length of premen-
tum and bearing only ten teeth; ligula narrow, less than one-third width of pre-
mentum at widest point and bearing 18 teeth and a dense brush of piliform setae
on anterior margin (Fig. 13). Lateral spines present on abdominal segments seven
to nine, dorsal hooks well developed on segments two to nine, ante-apical tu-
bercles of epiproct located at three-fifths of distance from base to tip.

Variations.—Adult males: total lengths 45.4—50.2 (avg. 48.8) mm, abdominal
lengths 31.4—35.3 (avg. 34.1) mm, hind wing lengths 27.0—-28.6 (avg. 28.0) mm.

VOLUME 95, NUMBER 1 201

Adult females: total lengths 47.5—-51.0 (avg. 48.7) mm, abdominal lengths 33.9-35.8
(avg. 34.7) mm, hind wing lengths 25.8-31.2 (avg. 29.3) mm. Exuviae: total
length (both sexes) 24.0—28.0 (avg. 26.4) mm. Above measurements from pre-
served material. Live final instar nymphs 23-24 mm.

The form and coloration of all adult specimens is generally uniform except that
the brown wash adjacent to the dorsal crest of the synthorax is often absent. The
occiput of females displays some variation in shape of the posterior outline viewed
dorsally often showing a slight convexity or concavity that may be artifactual.
The occipital horns display great variability in thickness and position (see Figs.
3-5). In addition, there are often adventitious spinules isolated some distance
from the main horns (Fig. 3), as a row between the horns and/or as lateral out-
growths of the large paired horns (Fig. 4).

Relationships.—Relationships among species of Ophiogomphus have not been
discussed. Species in eastern North America are morphologically distinctive and
have highly sympatric distribution areas. Closely related species pairs are un-
common in Ophiogomphus unlike many other genera and subgenera of Gomphi-
dae (Louton 1981). Three species of the genus share structural similarities that
seem to indicate a close relationship. Ophiogomphus bouchardi, O. carolinus,
and O. mainensis share: 1. similarly structured anterior occipital horns of the
female (though those of O. mainensis are adjacent), 2. rudimentary posterior
occipital horns and 3. terminal abdominal appendages of similar structure (though
exaggerated in O. mainensis). The close relationship of these three species is
further suggested by the allopatric nature of their distributional areas.

The simple matrix below facilitates the separation of the three above mentioned
similar species.

O. bouchardi O. carolinus O. mainensis
Yellow tibial stripe no! yes no
Middorsal thoracic stripe no yes yes

1 Proximal yellow spot about one-fifth length of tibia.

Nymphal habitat.—Nymphs of Ophiogomphus bouchardi were found in cherty
limestone gravel pockets in a small second order tributary (Will Hall Creek) of
the Harpeth River (Cumberland River System). The stream is predominantly
bedrock with gravel and sand deposits limited to fissures and pockets in pool
areas. Nymphs were also located in sand trapped by the roots of sedges growing
where bedrock fissures intersected the streambanks. The most common odonate
associates in this habitat were Stylogomphus albistylus, Boyeria vinosa and, less
commonly, Gomphus lividus. All three associates are widespread lotic general-
ists.

Acknowledgments

I would like to thank Christine Eason Louton and John L. Harris for assistance
in collecting nymphal material, Leah R. Baker for typing the manuscript, and Dr.
David A. Etnier for critically reviewing the manuscript.

202 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Literature Cited

Louton, J. A. 1981. Lotic dragonfly (Anisoptera: Odonata) larvae of the southeastern United States:
identification, distribution, and a consideration of the historical biogeography of the groups.
Unpublished dissertation, University of Tennessee, Knoxville.

Department of Zoology, University of Tennessee, Knoxville, Tennessee 37916.

Note added to galley proof: After this paper had gone to press, Dr. Minter
Westfall (University of Florida, Gainesville) informed me that the forthcoming
(December, 1981) issue of Odonatologica contained a description of a new spe-
cies of Ophiogomphus that would be conspecific with the one that I was de-
scribing (Carle, F. L. 1981. A new species of Ophiogomphus from eastern North
America, with a key to the regional species. Odonatologica 10(4):271—278).
Mr. Carle submitted his paper October 14, 1981, with the knowledge that I had
submitted a description of this same species a month earlier.

PROC. BIOL. SOC. WASH.
95(1), 1982, pp. 203-209

TWO NEW SPECIES OF ONUPHIS (ONUPHIDAE:
POLYCHAETA) FROM URUGUAY

Kristian Fauchald

Abstract.—Two new species of Onuphis (Onuphis) from Uruguay, O. (O.)
difficilis and O. (O.) orensanzi are described, based on material identified as
Onuphis setosa Kinberg by Orensanz (1974).

Kinberg (1865) described several species of Onuphis from the southern Atlantic
Ocean collected during the EUGENIE expedition. The material of most of these
species is in Riksmuseet, Stockholm, Sweden, with the exception of the material
of Onuphis setosa which has been lost (Roy Oleroed, in litt.).

The specimens of O. setosa must have been lost before 1930 since Augener
(1931:297) remarked that they were absent when he reviewed Kinberg’s collection
of onuphids from the La Plata region. Orensanz (1974:88) applied the name to a
species found in the type-area. Some discrepancies between Kinberg’s (1865,
1910) description and illustrations and those made by Orensanz suggested that it
was necessary to review Orensanz’ material. Three samples were made available
by the Director, M.N.H.N., Montevideo, Uruguay. As suspected, these speci-
mens cannot belong to Onuphis setosa Kinberg and are described below as two
new species.

The generic definitions and terminology are as in Fauchald (1980).

Onuphis (Onuphis) difficilis, new species
Fig. 1, Table 1

Onuphis setosa.—Orensanz, 1974:89, in part (not Kinberg, 1865:560).

Material.—Atlantic Ocean off Uruguay. 34°51'S, 54°04.5'W, 39 m depth, 14
April 1965, coll. A. Knipovich, st. 263 (holotype, M.N.H.N., Montevideo, I 1382,
| paratype, M.N.H.N., | paratype, USNM 69917).

Description.—The holotype is an incomplete specimen with 50 setigers that is
16.5 mm long and 1.7 mm wide with parapodia. It is pale buff-colored and lacks
distinct color patterns. Eyes are absent.

The prostomium (Fig. la) is a rounded lobe; the frontal palps are short and
triangular and the short peristomial cirri barely reach the middle of the prosto-
mium. The outer lateral occipital antennae reach setiger 1, the inner lateral an-
tennae reach setiger 7, and the median antenna reaches setiger 4. The occipital
ceratophores have up to 7 distinct rings on the lower half. Each ring is very
narrow and the rings are crowded near the base of the antennae making them
difficult to observe.

Branchiae are first present from setiger 6; all branchiae are branched; the first
branchia has 3 filaments; where best developed, posterior to setiger 21, each
branchia has 6 filaments. In one of the paratypes (USNM 69917), which consists
of 200 setigers, the last 50 setigers have reduced branchiae with only 3 or 4 fila-
ments.

204 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. |. Onuphis (Onuphis) difficilis holotype, N.M.H.N., Montevideo, I 1382: a, Anterior end,
dorsal view; b, Pectinate seta, median parapodium; c, Third parapodium, anterior view; d, Large
hook, third parapodium; e, Pseudocompound hook, third parapodium; f, Eighth parapodium, anterior
view; g, Anterior end, lateral view.

Ventral cirri (Fig. 1g) are cirriform in the first 8 setigers and postsetal lobes are
distinctly digitiform in the first 17 setigers in the holotype and in the first 11 or
23 setigers in the 2 paratypes. The first parapodia are no longer than those in the
second or third setiger (Fig. lc). In each of these parapodia the acicular lobe is
distally rounded, the presetal lobe follows the outline of the acicular lobe closely
except on the lower side where it is cut away from the base of the setae. A
distinct contraction fold is present across the middle of the frontal face. The
ventral cirrus and postsetal lobes are both triangular and of the same size. The
dorsal cirrus is more distinctly digitiform and twice as long as the ventral cirrus.
The ventral cirri and postsetal lobes become increasingly tapered posteriorly (Fig.

VOLUME 95, NUMBER 1 | 205

If). The dorsal cirrus remains of about the same length in all setigers but becomes
increasingly slender in posterior setigers.

Limbate and pectinate setae, pseudocompound, large and subacicular hooks
are present. Limbate setae are present in all setigers but are especially numerous
in the anterior third of the body. In the first 4 or 5 setigers they are short, slender,
and limited to the superior part of the fascicles. Each of the median and posterior
parapodia has 2 pectinate setae; each (Fig. 1b) is distinctly oblique distally and
has about 14 or 15 coarse teeth. The distal edges of the pectinate setae are curved.
Tridentate pseudocompound hooks (Fig. le) are present in the first 5 setigers; the
distal tooth is extremely long and slender and projects well beyond the edge of
the hood and the median and proximal teeth are short and slender. Large, tri-
dentate hooks (Fig. 1d) are present in setigers 3-8; each has a long, relatively
slender distal tooth, the median tooth is blunt and conical, and the proximal tooth
is slender and narrow. Bidentate subacicular hooks are first present in setiger 18
in the holotype and in setigers 15 and 18 in the 2 paratypes.

The maxillary formula (observed in the paratype in USNM 69917) is 1+1, 9+9,
11+0, 9+8, and 1+1.

Fragments of a ragged tube were present in the vial with the 3 specimens. The
inner lining of the tube appears to be soft, but with considerable tensile strength;
it is covered externally with loose debris in no recognizable pattern.

Kinberg’s illustrations and descriptions of Onuphis setosa show a species with
cirriform ventral cirri on less than 5 setigers, branchiae starting between setigers
10 and 20. The pseudocompound hooks are distally shown as bidentate and with
short, blunt hoods. Orensanz’ material from Uruguay have ventral cirri in at least
6 setigers, branchiae are present from setiger 6 and the pseudocompound hooks
are tridentate. The Uruguayan specimens resemble O. setosa basically in having
the median occipital antenna clearly shorter than the inner lateral ones, but this
is certainly not a unique character for this species.

Onuphis difficilis and O. orensanzi (see below) both resemble the same complex
of species, including O. fragilis Kinberg (1865:561; see also Fauchald 1980:808),
O. pulchra Fauchald (1980:814), O. simoni Santos, Day and Rice (1981:663), O.
vermillionensis Fauchald (1968:41) and O. virgata Fauchald (1980:819) in that all
these species have exclusively tridentate pseudocompound hooks, branched
branchiae starting at setigers 5-10 and ventral cirri present in at least 5 setigers.
A comparison of these species is given in Table 1.

Etymology.—The problems in differentiating species in this group prompted
the name.

Distribution.—Onuphis difficilis is known from one locality in shelf depths off
Uruguay in the southwestern Atlantic Ocean.

Onuphis (Onuphis) orensanzi, new species
Fig. 2, Tables 1 and 2

Onuphis setosa.—Orensanz, 1974:94, in part (not Kinberg, 1865:560).

Material.—Atlantic Ocean off Uruguay, 34°51'S, 52°35’W, 83 m, 12 April 1965,
coll. A. Knipovich, st. 250 (holotype, M.N.H.N., Montevideo, I 1380, 129 para-
types, M.N.H.N., USNM 69918). Atlantic Ocean off Uruguay, 35°20.8’S,

206 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Comparison of selected species of Onuphis (Onuphis). The columns are: 1, Number of
rings in the occipital ceratophores; 2, Number of segments with cirriform ventral cirri; 3, Number of
segments with pseudocompound hooks; 4, Last segment with large hooks; 5, First branchia present;
6, Maximal number of branchial filaments; 7, Number of teeth in the pectinate setae; 8, First segment
with subacicular hooks.

Name 1 2 3 4 5 6 7 8
difficilis 7-8 qo 5 8 6 6 15 18
fragilis 0 7 a 12 6 D) v 25
orensanzi 3-4 5 4 6 6 3 10 13
pulchra 4-5 9 6 15-19 6 4 50 16-20
simoni 3 6-8 5) 16 6-9 7-8 16-19 7
vermillionensis 4-5 9 8 10 7 4 14 12
virgata 4-5 ails 7 DIL, S=]) 5 8 Mpls)

52°50.4’W, 130 m, 14 April 1965, coll. A. Knipovich, st. 260 (15 paratypes,
M.N.H.N., Montevideo, I 1383).

Description.—The holotype is a complete specimen with 153 setigers that is
33.15 mm long and 2.50 mm wide with parapodia. The body is cylindrical and
pale pink in color; distinct color patterns are absent. The prostomium is a short
rounded lobe with the occipital antennae attached medially. The frontal palps are
short and slender. At the posterior margin of the prostomium is a small, distinct
tubercle. The outer lateral occipital antennae reach setiger 2, the inner lateral
antennae reach setiger 5 and the median antenna reaches setiger 3. The inner
lateral ceratophores have up to 4 basal rings. The peristomial antennae are short
and slender and do not reach beyond the base of the occipital antennae.

Branchiae are present from setiger 6 to setiger 92 in the holotype. The first 15
and the last 30 pairs of branchiae are single filaments; all other branchiae have
either 2 or 3 filaments where best developed.

The first parapodia are all similar in size; the acicular lobes are rounded and
the presetal lobe follows the outline of the acicular lobe closely. The postsetal
lobes and the ventral cirri are of the same length in the first parapodium (distorted
in Fig. 2a), in the second and third parapodia the ventral cirri are distinctly longer
than the postsetal lobe. The ventral cirri are digitiform; the postsetal lobes are
triangular. The dorsal cirri are longer than the ventral cirri in all setigers; each
is digitiform but becomes slender and thread-like in posterior setigers. Ventral
cirri are cirriform in the first 5 setigers in all specimens. Postsetal lobes are
digitiform in the first 10 setigers.

Limbate and pectinate setae, pseudocompound, large and subacicular hooks
are present. Limbate setae are present in all setigers, but are especially common
in the first third of the body. Distally oblique pectinate setae with about 10 teeth
each are present in median and posterior setigers. Usually 2 pectinate setae are
present in a parapodium. Tridentate pseudocompound hooks with short blunt
hoods are present in the first 3—S setigers; the distal tooth is only slightly longer
than the median and proximal teeth and does not project beyond the hood. Large
hooks are present from setiger 3 to about 6; each large hook is tridentate with 2
large, strongly curved teeth basally and a short, slender tooth distally. The distal

VOLUME 95, NUMBER 1 207

\ 0.00Imm_c.d,e
SZ A
0.1mm b, f
t U

lmm a,g

Fig. 2. Onuphis (Onuphis) orensanzi: a, c—e and g, holotype M.N.H.N., Montevideo, I 1380, b and
f, paratype USNM 69918. a, Anterior end, lateral view; b, Seventh parapodium, anterior view; c,

Pectinate seta, median parapodium; d, Large hook, third parapodium; e, Pseudocompound hook,
third parapodium; f, Third parapodium, anterior view; g, Anterior end, dorsal view.

tooth is often broken, but the base of the tooth can always be recognized. Bi-
dentate subacicular hooks are present from setiger 12 or 13.

The maxillary formula (investigated in 3 paratypes from USNM 69918) is 1+1,
oo, / +0, 4+6—7, and 1+].

208 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Survey statistics for Onuphis (Onuphis) orensanzi.

Range Mean s.d. V. N

Occipital antennae

Outer lateral reach number 1-2 1.03 0.16 0.03 39

Inner lateral reach number 3-9 D278) 1.03 1.06 35)

Median reaches number 2-4 3.19 0.66 0.44 37)

Number of rings 3-4 3.78 0.42 0.18 40
Branchiae

From setiger number 6 invariant

To setiger number 76-92 87.00 5.60 31.33 7

Number of filaments 1-3 De) 0.60 0.36 40
Cirriform ventral cirri to setiger num-

ber 3 invariant
Digitiform postsetal lobes to setiger

number 8-10 9.83 0.50 0.25 40
Pseudocompound hooks to setiger

number 3=5 3.90 0.44 0.19 40
Number teeth in pseudocompound hooks 3 invariant
Subacicular hooks first present from

number 10-14 12.53 0.82 0.67 40
Volume through setiger 10 0.03—2.68 1.05 0.54 0.29 40

Mean length per setiger (x 100): 18.10
Mean length per setiger (x 100), st. 250: 16.12
Mean length per setiger (< 100), st. 260: 22.74

Tubes are slender, with a thin inner lining and thin cover of sediment particles.

All specimens from station 260 were investigated in detail, as were 25 of the
130 specimens from station 250. The remaining 105 specimens were only cursorily
examined, but do not appear to differ in the distribution of ventral cirri, branchiae,
postsetal lobes, or in the number of rings on the ceratophores.

The differentiation of O. orensanzi and related species is indicated in Table 1.

Etymology.—The species is named in honor of the Argentinian polychaetologist
José Maria Orensanz whose papers on the southwest Atlantic fauna have added
considerably to our knowledge of the polychaetes.

Distribution.—O. orensanzi is known from two localities off the coast of Uru-
guay and may be widespread in the southwest Atlantic Ocean.

Acknowledgments

I would like to thank Dr. José Maria Orensanz for his efforts in arranging the
loan of the specimens housed at the Museo Nacional de Historia Natural, Mon-
tevideo, and the Director of that Institution, Dr. Miguel A. Klappenbach for
allowing me the loan of the material. My colleague, Dr. Meredith L. Jones, kindly
read and commented on the manuscript.

Literature Cited

Augener, H. 1931. Die bodensassigen Polychaeten nebst einer Hirudinee der Meteor-Fahrt.—Mittei-
lungen aus dem Hamburgischen Zoologische Staatsinstitut und Museum 44:279-313.

VOLUME 95, NUMBER 1 209

Fauchald, K. 1968. Onuphidae (Polychaeta) from western Mexico.—Allan Hancock Monographs in

Marine Biology 3:1-82.

. 1980. Onuphidae (Polychaeta) from Belize, Central America, with notes on related taxa.—

Proceedings of the Biological Society of Washington 93(3):797-829.

Kinberg, J. G. H. 1865. Annulata nova.—Oefversigt af Kungliga Vetenskaps-Akademiens For-

handlingar, Stockholm, 1864 (vol. 21):559-574.

. 1910. Annulater. In: Kungliga Svenska Fregatten Eugenie’s Resa omkring jorden under befal

af C. A. Virgin aren 1851—1853.—Vetenskapliga lagttagelser pa Konung Oscar den Forstes

Befallning utgifna delen. Zoologi 3:33—78 (issued posthumously by Hajalmar Theel).

Orensanz, J. M. 1974. Los anelidos poliquetos de la provincia biogeografica Argentina V. Onuph-
idae.—Physis, Buenos Aires, Seccion A 33(86):75—122.

Santos, S. L., R. Day, and S. A. Rice. 1981. Onuphis simoni, a new species of polychaete (Poly-
chaeta: Onuphidae) from south Florida.—Proceedings of the Biological Society of Washington
94(3):663—668.

Department of Invertebrate Zoology, Smithsonian Institution, Washington,
D.C. 20560.

PROC. BIOL. SOC. WASH.
95(1), 1982, pp. 210-221

REMARKS ON THE STYLASTERINE FAUNA OF THE
WEST INDIES, WITH THE DESCRIPTION OF
STYLASTER ANTILLARUM, A NEW SPECIES

FROM THE LESSER ANTILLES
(CNIDARIA: HYDROZOA:
STYLASTERINA)

Helmut Zibrowius and Stephen D. Cairns

Abstract.—Stylaster antillarum, a new species of stylasterine coral from the
Lesser Antilles, is described and illustrated. The 23 species of Stylasterina now
known from the West Indies are listed and briefly discussed. Inaccurate distri-
butional patterns of some of these species, usually resulting from incorrect iden-
tifications, are noted.

Introduction

Most of the information on the Stylasterina of the West Indies (between Flor-
ida, Central America and South America), comprising descriptions of species,
nomenclatural changes, additional records and inventories, can be found in the
following publications: Pallas (1766), Duchassaing and Michelotti (1861, 1864),
Pourtales (1867, 1868, 1871, 1874, 1878), Saville Kent (1870, 1871), Lindstrom
(1877), Moseley (1879, 1881), Broch (1936, 1942), Boschma (1951, 1955, 1957,
1962, 1964a, 1964b, 1964c, 1965), Squires (1962), Roos (1971), Vervoort & Zi-
browius (1981) and Zibrowius (in press).

Until now, stylasterine corals have been reported from the West Indies under
28 specific and one subspecific names which are listed below. Six of these species
and the one subspecies are considered to be junior synonyms or dubious records,
resulting in 23 valid species records (including the new species) for the area. The
invalid and dubious species records are marked by an asterisk. If not otherwise
indicated, the original descriptions of these species are based on material from the
West Indies. :

Stylaster complanatus Pourtalés, 1867

Stylaster duchassaingi Pourtales, 1867
(new name for Stylaster elegans Duchassaing & Michelotti, 1864)

Stylaster echinatus Broch, 1936

Stylaster antillarum, new species

*Stylaster elegans Duchassaing & Michelotti, 1864
(Gunior homonym of Stylaster elegans Verrill, 1864, a species from the Marshall Islands, Pa-
cific; renamed S. duchassaingi by Pourtalés, 1867)

Stylaster erubescens Pourtaleés, 1871

*Stylaster eximius Saville Kent, 1871
(new name for Stylaster elegans Duchassaing & Michelotti, 1864; junior synonym of S. du-
schassaingi)

*Stylaster eximius forma atlanticus Broch, 1936
(Gunior synonym of S. duchassaingi)

Stylaster filogranus Pourtales, 1871

Stylaster punctatus Pourtales, 1871

VOLUME 95, NUMBER 1 211

Stylaster roseus (Pallas, 1766)
(originally described as Madrepora)
*Stylaster sanguineus Milne Edwards & Haime, 1849
(type-locality Australia; dubious record for western Atlantic: see text)
Allopora miniata Pourtalés, 1868
Stenohelia challengeri Boschma, 1951
(new name for Stenohelia profunda Moseley, 1881)
*Stenohelia maderensis (Johnson, 1862)
(originally described as Allopora, type-locality Madeira, East Atlantic; dubious record for
western Atlantic: see text)
Stenohelia virginis (Lindstrom, 1877)
(originally described as Crypthelia)
Crypthelia peircei Pourtales, 1867
*Crypthelia pudica Milne Edwards & Haime, 1849
(type-locality Philippine Islands; dubious record for western Atlantic)
Distichopora barbadensis Pourtalés, 1874
Distichopora cervina Pourtales, 1871
Distichopora contorta Pourtales, 1878
Distichopora foliacea Pourtalés, 1868
Distichopora sulcata Pourtales, 1867
*Errina (Errina) aspera (Linné, 1767)
(originally described as Millepora, type-locality Mediterranean; not present in western Atlantic:
misidentification of Broch, 1942)
Errina (Lepidopora) carinata Pourtales, 1867
(originally described as Heliopora)
Errina (Lepidopora) cochleata Pourtales, 1867
Errina (Lepidopora) decipiens Boschma, 1964
Errina (Lepidopora) glabra Pourtales, 1867
Pliobothrus symmetricus Pourtalés, 1868
Pliobothrus tubulatus (Pourtales, 1867)
(originally described as Heliopora)

General remarks.—A review of the literature and a preliminary study of nu-
merous specimens in various museum collections (National Museum of Natural
History, Washington, D.C.; Yale Peabody Museum; Museum of Comparative
Zoology; Muséum National d’Histoire Naturelle, Paris; Rijksmuseum van Na-
tuurlike Historie, Leiden; Zoologisch Museum, Amsterdam; British Museum
(Natural History), London; Royal Scottish Museum, Edinburgh; Zoologisk Mu-
seum, Copenhagen; Naturhistoriska Riksmuseet, Stockholm) provide the bases
for the following remarks on the synonymy and the distribution of the West Indian
Stylasterina; however, a monographic study including a revision of all previous
records is greatly needed.

Stylaster elegans (name preoccupied), Stylaster eximius, S. eximius forma at-
lantica and Stylaster duchassaingi are all names given to the same species for
which the latter name must be retained.

Stylaster complanatus has characters somewhat intermediate between typical
Stylaster and Stenohelia. Here it is listed with the former genus although some
authors referred it to Stenohelia (Saville Kent 1870; Broch 1936; Boschma 1957,
1964b,c). Broch (1936) incorrectly synonymized it with Stenohelia virginis.

The shallow-water specimens of Stylaster punctatus (which Pourtalés seemed
to consider as the typical form) are identical with Stylaster roseus.

The new name Stenohelia challengeri had been introduced by Boschma (1951)
for Stenohelia profunda Moseley, 1879, because of homonymy with Allopora

212 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

profunda Moseley, 1879, in case the closely related genera Stylaster, Allopora
and Stenohelia were synonymized, as has been suggested previously by some
authors. However, Boschma’s later papers considered these as separate genera
(a point of view now generally accepted), and therefore there was no real need
for that new name.

Errina (Lepidopora) cochleata has been incorrectly described as a bryozoan,
Hornera galeata Smitt, 1872 (see Zibrowius, in press).

Most of the West Indian species appear to be endemic to that area, or at least
to a larger area in the West Atlantic, whereas only a few species are amphi-
Atlantic. The occurrence in the eastern Atlantic of Stylaster erubescens (off west-
ern Brittany, between Hebrides and Faeroe, between Orkney and Faeroe, off
southeast Iceland) and of Pliobothrus symmetricus (Hyeéres Banc, Azores, off
western Brittany, off western Ireland, Faeroe, Norway, off southeast Iceland) is
confirmed. In addition, a still unnamed species of Crypthelia known from the
eastern Atlantic (Hyeres Banc, Azores, Madeira) has been recognized among
Pourtalés material from the Lesser Antilles (Museum of Comparative Zoology).

Not all indications in the literature on amphi-Atlantic distributions are correct.
Errina (Errina) aspera has been reported by Broch (1942) from the West Indies
(detailed origin unknown) based on a small fragment of pink Errina (Naturhis-
toriska Riksmuseet, Stockholm, No. 45). However, this is not E. aspera, oth-
erwise known from the southwest Mediterranean and the eastern Atlantic, but
probably an Antarctic or Subantarctic species (mislabelled), as already suggested
by Boschma (1965:2). Nothing similar of authentic West Indian origin has yet
been found in the various museum collections.

The case of Stenohelia maderensis, reported by Boschma (1964b, 1964c) from
the West Indies (St. Vincent), needs further investigation.

Indications in the literature of species of even wider distribution, occurring in
both the West Indies and in the Indo-Pacific, are also doubtful and are possibly
all caused by mislabelling (origin confused), inversion of labels in the collections,
or misidentification. This is sufficiently confirmed in some cases. For instance,
Stylaster erubescens (?) sensu Moseley, 1881, from the Kermadec Islands, is a
Conopora (British Museum (Nat. Hist.) 1880.11.25.178).

Typical specimens of Stylaster sanguineus, a pink shallow-water species well
known from the central and southwest Pacific, are labelled as being from Florida
in the Museum of Comparative Zoology and the Yale Peabody Museum. They
may well be those mentioned by Pourtalés (1871:83) as undistinguishable from
authentic Pacific specimens. A label-inversion error with specimens of the pink
shallow-water species Stylaster roseus from the West Indies, might best explain
this unusual, still unverified distribution. Furthermore, S. roseus is mentioned on
some of the old labels, either as the original identification or as a synonym of S.
sanguineus. Boschma (1957) reported the reverse case of Stylaster roseus oc-
casionally being mentioned from the Pacific (Hawaii, Samoa).

Boschma (1957) further points out that material from the Loyalty Islands used
in an anatomical study (England 1926) had erroneously been cited as Stylaster
filogranus, the latter being a West Indian species.

A detailed study will probably confirm that the Stylaster from Mauritius, In-
donesia, the Philippine Islands and the Sea of Okhotsk referred to as S. eximius
(see Boschma 1957) are not only ‘‘forma’’ or “‘facies’’ but species distinct from

VOLUME 95, NUMBER 1 213

the West Indian one, for which the name Stylaster duchassaingi has priority over
S. eximius. A similar result can reasonably be expected from the comparison of
Stenohelia virginis (Lindstrom, 1877) with Stylaster virginis sensu Hickson and
England, 1905, from Indonesia.

Boschma (1964c) affirms that Moseley’s (1881) two records of Stenohelia pro-
funda (renamed Stenohelia challengeri) from the West Indies (type-locality) and
the Kermadec Islands are indeed the same species. If he is correct, it would be
a very unusual distribution, but it should be remembered that specimens of var-
ious zoological groups in the “‘Challenger’’ collections have confused origins. On
the other hand, specimens from the Galapagos Islands originally reported as
Stenohelia profunda by Marenzeller (1904) have been described by Boschma
(1964b) as a distinct species, Stenohelia robusta.

World-wide, several species have probably been identified as Crypthelia pu-
dica. Moseley’s (1881) record of C. pudica from the West Indies (Sombrero) is
considered with suspicion; until more detailed comparisons are made, we do not
assume that it occurs both in the western Pacific (type-locality: Phillipine Islands)
and in the western Atlantic.

A new species of Stylaster, distinct from all those previously reported from the
West Indies, is described below. The unstudied material from the cruise of the
United States Coast Survey steamer “‘Blake’’ in 1878/79 to the Caribbean Islands
(eight stations) was discovered in the collection of the Museum of Comparative
Zoology during a visit in summer, 1980. The original labels printed for that cruise
read: ‘U.S. Coast Survey, C. P. Patterson, Supt.-Caribbean Islands exploration.
U.S.C.S., S. Blake, Alex. Agassiz, 1878—79.’’ They are marked in pencil with the
station number, depth (in fathoms), and the name of the nearest island. Comple-
mentary information on the stations was obtained from Smith’s compilation
(1889). No preliminary identification, generic or specific, was found with the
specimens. Pourtales’ (1880) report on this cruise of 1878/79 deals only with the
Scleractinia and Antipatharia. He probably intended to publish a separate report
on the Stylasterina, whereas his previous papers include both Scleractinia and
Stylasterina.

Stylaster antillarum, new species
Figs. 14

Types.—The nearly complete colony from ‘‘Blake’’ Sta. 241 is designated as
holotype. The other colonies, branches, and fragments from ‘“‘Blake’’ stations
213, 215, 216, 218, 219, 231, and 238 are designated as paratypes. Depository:
holotype and most paratypes at MCZ; two paratypes at USNM (‘‘Blake”’ sta.
216, USNM 60350; ‘‘Blake’’ sta. 219, USNM 60349).

Description.—The **Blake’’ material is in good condition; all had been collected
alive and then dried. For the present study it has been cleaned of the dry tissue
and centenary dust with sodium hypochlorite solution.

The holotype is a regularly flabellate and moderately branched colony rising
from a strong, encrusting base that is 7.5 mm wide. About 40 mm high and 39
mm wide, the holotype is the largest of the specimens from the ‘‘Blake”’ stations.
Most of the other specimens are just branches or fragments and considerably
smaller. Colonies comprising the entire base or at least the basal part of the trunk
are rare. The coenosteum bears distinct, discontinuous carinae, which are most

214 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Stylaster antillarum type-material from “*‘Blake’’ cruise 1878/79.

Station Date Area Coordinates Depth Quantity

213. «12.22.1879 Martinique 14°32'38"N, 61°06'40"W 357 fm—653 m 1 branch

21Se MS 221879 eS t. lence 13°51'30"N, 61°03'45’"W 226fm—414 m 2 branches

26: AS 2879S twleueia 13°51’45"N, 61°03’30"W *153 fm—280 m 15 colonies, branches,
fragments

2ANSe NSOSTOe eet eucta 13°49'12’N, 61°04'40"W__ 164 fm-—300 m 1 colony, | branch

21D AS 2879 Ste acta 13°49’50"N, 61°03’50"W 151 fm—276 m 9 colonies, branches,
fragments

231 20.22.1879) St) Vincent “132710 Ne ol 17 1s WwW 95; tim— 17/4 om -1 branch

238 23.2.1879 Grenadines: 12°46'10"N, 61°23'35”W *126 fm—231 m 1 branch
Cannuan

241 24.2.1879 Grenadines: 12°28'22"N, 61°32’18"W 163 fm—298 M 1 colony
Carriacou

* The original labels indicate slightly different depths, respectively 154 fathoms and 127 fathoms.

well developed near a cyclosystem, one carina corresponding to each dactylo-
pore. The carinae are about 32-37 wm wide, about 64 um tall, and variable in
length, ranging from 0.1 mm to several millimeters. The distance between two
adjacent carinae varies between 0.20-0.30 mm. The coenosteum lying between
the carinae is usually slightly concave and is penetrated by coenosteal slits which
are also aligned parallel to the carinae. The slits are variable in length, ranging
from 20-80 um, but are consistently 8-9 wm wide. Distinct, rounded coenosteal
granules are not present; however, the coenosteal surface is covered by irregular
deposits of approximately 3-6 x 2-3 wm in size, particularly noticeable at the
edges of coenosteal slits (Fig. 2D).

Branching is strictly in one plane as the result of simultaneous budding to the
left and to the right from below the top part of older cyclosystems. Otherwise,
branches lengthen by alternate budding to the left and right. Angle of budding
about 60—70°. This sympodial pattern persists even on the main branches. Cyclo-
systems occur only on the lateral branch edges; the anterior and posterior sides
of the colony are entirely devoid of cyclosystems. |

Well-developed young distal and peripheral cyclosystems are long, cylindrical
structures, circular in cross section, except distally, where they are compressed
in the flabellar plane. They are about 3.2—3.6 mm high and 0.9 mm in diameter
at the base. The thickness of the wall of a cyclosystem is about 0.2 mm, and the
lesser axis of the distal part of a cyclosystem is about 0.9-1.0 mm, the greater
axis, 1.2-1.3 mm. Older cyclosystems are not much taller but do become pro-
nouncedly thicker (thickening of the wall), becoming proportionally more com-
pressed at the top (lesser axis 1.0 mm, greater axis up to 1.6 mm). Initial stages
of new cyclosystems start as low annular ridges, 0.7 mm wide, below the tops of
older ones.

Seen from the anterior and posterior sides of the colony, the top of the cyclo-
system is distinctly flabellate with a rounded outline, the pseudosepta and dac-
tylopore openings near the greater (longitudinal) axis being much lower than those
near the lesser (transverse) axis. Adjacent to the greater axis the dactylopores

VOLUME 95, NUMBER 1 215

Fig. 1. Types of S. antillarum: A, Holotype colony from ‘‘Blake’’ sta. 241, MCZ, x2.0; B,
Branchlet of holotype with prominent ampullae and circular depressions left by ruptured ampullae,
x6.3; C, Detail of paratype colony from ‘‘Blake’’ sta. 216, MCZ, with ampullae occurring singly or
in pairs below the flared top of the cyclosystems, 6.3; D, paratype colony from ‘‘Blake’’ sta. 219,
MCZ, with large acrothoracic cirriped crypt in the fractured base, <4.3.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

216

, USNM 60350: A-B, Lateral and oblique

‘‘Blake’’ sta. 216

Paratypes of S. antillarum from

nicus
views of cyclosystems and coenosteal ridges,

one ridge corresponding to each dactylopore, x44,

x 640; E, Oral view of cyclosystem

b]

, Coenosteal slit

x106; D
with 15 dactylopores, x50, SEM stereomicrographs.

’

respectively; C, Coenosteal texture

’

x34

VOLUME 95, NUMBER 1 ZAI

Fig. 3. Paratypes of S. antillarum: A, ‘‘Blake’’ sta. 216, longitudinal section of gastropore re-
vealing gastrostyle, x24; B, Gastrostyle of figure 3A, 110; C, ‘‘Blake’’ sta. 219, USNM 60349
gastrostyle and inner ring of granules, X110, SEM stereomicrographs; D, ‘‘Blake’’ sta. 216, dacty-
lostyle, X680; E, Gastrostyle tip of fig. 2C, x340.

218 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

are closely spaced, separated by very distinct lamellar pseudosepta; adjacent to
the lesser axis they are widely spaced, separated by thick pseudosepta which are
less prominent and more flattened toward the gastropore tube. Upper rim of
cyclosystem crenulate; upper part of outside surface carinate, with carinae cor-
responding to dactylopores and depressions to pseudosepta.

Dactylopores are deep, narrow pits parallel to the central gastropore tube,
separated from the latter by a thin wall except at the upper part where the narrow
slit-like dactylotomes permit open communication. The gastropore tube is cylin-
drical, up to 3 mm long, and about 0.40-0.43 mm in diameter. The basal 0.5 mm
of the tube, the gastrostyle chamber, is narrower, usually only 0.3 mm in diam-
eter. At the boundary of the gastrostyle chamber with the larger, upper part there
is a distinct ring of granules, the ring being about 0.2 mm thick. The granules are
thick and blunt, about 50 um tall and 35-40 um thick, and are closely spaced,
often fusing. This ring of granules further constricts the gastropore tube to a
diameter of about 0.20-0.25 mm.

Gastrostyles are conical to bullet-shaped, up to 0.55 mm tall and 0.20 mm
wide. Height : width ratios of those styles measured ranged from 2.67—2.90. The
style occupies the lower 15-19% of the gastropore tube, the delicate tip usually
extending just above the ring of blunt granules. Styles are highly sculptured with
spines which are particularly well developed at the tip where they measure up to
36 wm long and 8.5 wm wide (Fig. 3E). No ridges are present on the styles.

Dactylostyles are very poorly developed, composed of small, irregular, angular
deposits measuring about I5 x 5 wm. These deposits are usually fused in a line
forming the dactylostyle.

The number of dactylopores varies from 10 to 18 in the 200 cyclosystems
examined in detail; 14 dactylopores per cyclosystem is both the median and the
mode, 14.35 is the average.

Dactylopores per cyclosystem 10 1] 12 13 14 15 16 17 18
Number of cyclosystems 7 9 16 28 44 40 31 17 8

The ampullae are prominent outgrowths on both the anterior and posterior
sides of the colony, located just below the flared top of the cyclosystem, at the
same level from which the younger cyclosystems bud. They measure up to 0.57
mm tall and 0.61 mm broad. Ampullae occur singly or in pairs and the same
cyclosystem may have ampullae on both its anterior and posterior sides. Their
roughly hemispherical shapes are obscured by irregular crests and small knobs.
Some ampullae have a tiny pore near the base, probably an efferent duct. Rather
frequently ampullae are ruptured, leaving a circular depression.

Remarks.—Cavities caused by acrothoracic cirripeds are present in the base
of two colonies, one from ‘““Blake’’ sta. 216, the other from ““Blake”’ sta. 219.
No other symbionts are known from Stylaster antillarum.

Discussion.—The combination of characters described above distinguishes S.
antillarum from all previously described species of Stylaster (see Boschma 1957;
Vervoort & Zibrowius 1981). As far as could be deduced from the literature, only
Stylaster multiplex Hickson & England, 1905, and Stylaster crassior Broch, 1936,
show some resemblance to S. antillarum, mainly by the somewhat elongate tops
of their cyclosystems. However, comparison with S. multiplex (type-material
from Indonesia) and S. crassior (from the Comoros, “‘Le Suroit’’ cruise, Ben-

VOLUME 95, NUMBER 1 219

Fig. 4. Paratypes of S. antillarum from ‘‘Blake’’ sta. 216, USNM 60350: A, Cross section of
gastropore tube just above ring of granules; gastrostyle tip in center; rudimentary dactylopore tubes
encircle gastropore, x50; B, Two fused granules from the ring of granules of fig. 4A, <x680; C,
cyclosystem flanked by irregularly shaped ampullae, x30.

thedi, 1977; the type-material from Mauritius was not available) indicates that
these forms, similar to each other, differ considerably from the West Indian
species. The tops of their cyclosystems are not as prominent or flabellate as in
S. antillarum, and their dactylopores and pseudosepta are more uniformly
spaced. Other differences concern the depth of the gastropore tube, the surface
structure of the branches, and the ampullae.

Acknowledgments

We would like to thank Dr. Herbert W. Levi (Museum of Comparative Zool-
ogy, Harvard University) for making available to us the specimens reported in

220 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

this paper and for the cooperation extended toward the senior author during his
visit to the MCZ in 1980.

Literature Cited

Boschma, H. 1951. Notes on Stylasterina (Hydrocorallia).—-Proceedings Koninklijke Nederlandse

Akademie van Wetenschappen (C)54(5):451-458.

. 1955. The specific characters of the coral Stylaster roseus.—Deep-Sea Research 3, Supple-

ment: 134-138. .

. 1957. List of the described species of the order Stylasterina.—Zoologische Verhandelingen,

Leiden 33:1-72.

. 1962. Notes on the Stylasterine coral Allopora miniata.—Proceedings Koninklijke Neder-

landse Akademie van Wetenschappen (C)65(3): 195-204, pl. 1-2.

. 1964a. Errina (Lepidopora) decipiens, a new Stylasterine coral from the West Indies.—

Proceedings Koninklijke Nederlandse Akademie van Wetenschappen (C)67(2):55-63, pl. 1.

. 1964b. On the Stylasterina of the genus Stenohelia.—Proceedings Koninklijke Nederlandse

Akademie van Wetenschappen (C):67(2):64—73, pl. 1-2.

——. 1964c. Further notes on the Stylasterine corals Stenohelia challengeri and Stenohelia mader-

ensis.—Proceedings Koninklijke Nederlandse Akademie Wetenschappen (C)67(2):78-84, pl.

|Z.

. 1965. Further notes on Stylaster roseus (Pallas). I & I1.—Proceedings Koninklijke Neder-

landse Akademie Wetenschappen (C)68(2):229-250, pl. 1-4.

Broch, H. 1936. Untersuchungen an Stylasteriden (Hydrokorallen). Teil 1.—Skrifter utgitt av Det

Norske Videnskaps-Academi i Oslo, I Mat.-Naturv. Klasse, 8:1—103, pl. 1-13.

. 1942. Investigations on Stylasteridae (Hydrocorals).—Skrifter utgitt avy Det Norske Viden-

skaps-Academi 1 Oslo, I Mat.-Naturv. Klasse, No. 3:1—113, pl. 1-6.

Duchassaing, P., and J. Michelotti. 1861. Mémoire sur les coralliaires des Antilles—Mémoires de

1’ Academie des Sciences de Turin (2)19:279-365, pl. 1-10.

. 1864. Supplément au mémoire sur les coralliaires des Antilles Mémoires de |’ Academie

des Sciences de Turin (2)23:97—206, pl. 1-11.

England, H. M., 1926. Development of the gonophores of the Stylasteridae.—Proceedings of the
Zoological Society of London (1926):265—283.

Hickson, S. J., and H. M. England. 1905. The Stylasterina of the Siboga expedition.—Siboga-Ex-
peditie 8:1—26, pl. 1-3.

Lindstrom, G. 1877. Contributions to the actinology of the Atlantic Ocean.—Kongliga Svenska Ve-
tenskaps-Akademiens Handlingar, Stockholm 14(2):1—26, pl. 1-3.

Marenzeller, E. von. 1904. Stein- und Hydro-korallen.—Bulletin of the Museum of Comparative
Zoology 43(2):75-87, pl. 1-3.

Moseley, H. N. 1879. On the structure of the Stylasteridae, a family of the Hydroid stony corals.—

Philosophical Transactions of the Royal Society of London 169, 1878:425-503, pl. 34-44.

. 1881. Report on certain Hydroid, Alcyonarian and Madreporarian corals procured during the

voyage of H.M.S. Challenger in the years 1873—1876.—Report on the Scientific Results of the

Voyage of H.M.S. Challenger during the years 1873-1876. Zoology 2:1—248, 32 pl.

Pallas, P. S. 1766. Elenchus zoophytorum . . . Hagae Comitum xvi-28-451 pp.

Pourtales, L. F. de. 1867. Contributions to the fauna of the Gulf Stream at great depths.—Bulletin

of the Museum of Comparative Zoology 1(6):103—120.

. 1868. Contributions to the fauna of the Gulf Stream at great depths (24 series).—Bulletin of

the Museum of Comparative Zoology 1(7):121—-142.

. 1871. Deep-sea corals—lIllustrated Catalogue of the Museum of Comparative Zoology 4: 1-93,

pl. 1-8.

. 1874. The zoological results of the Hassler Expedition. Deep-sea corals.—Illustrated Cata-

logue of the Museum of Comparative Zoology 8:33-49, pl. 6-9.

. 1878. Report on the results of dredging, under the supervision of Alexander Agassiz, in the

Gulf of Mexico, by the U.S. coast survey steamer “‘Blake.’’ Corals.—Bulletin of the Museum

of Comparative Zoology 5(9):197—212, pl. 1.

Roos, P. J. 1971. The shallow-water stony corals of the Netherlands Antilles.—Studies on the Fauna
of Curacao and other Caribbean Islands 37(130):1—108, pl. 1-53.

VOLUME 95, NUMBER 1 22)

Saville Kent, W. 1870. On a new genus of Madreporaria or stony corals (Stenohelia).—Annals and

Magazine of Natural History, London (4)5:120-123.

. 1871. On some new and little-known species of Madrepores, or stony corals, in the British

Museum collection.—Proceedings of the Zoological Society of London (1871):275-286, pl.

23-25.

Smith, S. 1889. Lists of the dredging stations of the U.S. Fish Commission, the U.S. Coast Survey,
and the British steamer Challenger, in North American waters, from 1867 to 1887, together
with those of the principal European government expeditions in the Atlantic and Arctic
Oceans.—United States Commission of Fish and Fisheries, Report for 1886:873—1017, maps.

Smitt, F. A. 1872. Floridan Bryozoa collected by Count L. F. de Pourtales. Part I.—Kungliga Svenska
Vetenskaps-akademiens Handlingar, Stockholm 10(11):1-20, pl. 1-5.

Squires, D. F. 1962. Additional Cretaceous and Tertiary corals from New Zealand.—Transactions
of the Royal Society of New Zealand 88(4):133-—150, 4 pls.

Vervoort, W., and H. Zibrowius. 1981. Annotations on H. Boschma’s work on Hydrocorals (Mil-
leporina, Axoporina, Stylasterina) with additions to his list of the described species of Stylas-
terina.—Zoologische Verhandelingen, Leiden 181:1—40.

Zibrowius, H. in press. Identification des pretendus Bryozoaires (“‘Hornera’’) de Smitt et de Calvet
a des Hydrocoralliaires Stylasterina.—Bulletin Muséum National d’ Histoire Naturelle, Paris.

(HZ) Station Marine d’Endoume, Marseille 13007, France; (SDC) Department
of Invertebrate Zoology, National Museum of Natural History, Smithsonian In-
stitution, Washington, D.C. 20560.

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CONTENTS

A new species of skate from Western Australia with comments on the status of Pavoraja —

Whitley, 1939 (Chondrichthyes: Rajiformes) John D. McEachran and Janice D. Fechhelm
A new species of Gymnure, Podogymnura, (Mammalia: Erinaceidae) from Dinagat Island,

Philippines Lawrence R. Heaney and Gary S. Morgan

Checklist of the fishes of the central and northern Appalachian Mountains
Jay R. Stauffer, Jr., Brooks M. Burr, Charles H. Hocutt, and Robert E. Jenkins
Two new genera of deep-sea Polychaete worms of the family Ampharetidae and the role of one
species in deep-sea ecosystems Robert Zottoli
The karyotype of the Eurasian flying squirrel, Pteromys volans (L.), with a consideration of

karyotypic and other distinctions in Glaucomys spp. (Rodentia: Sciuridae)

V. R. Rausch and R. L. Rausch
A new species of the genus Echininus (Mollusca: Littorinidae: Echinininae) with a review
of the subfamily . Joseph Rosewater
A new dwarf Sphaerodactylus from Haiti (Lacertilia: Gekkonidae) Richard Thomas
Amerigoniscus malheurensis, new species, from a cave in western Oregon (Crustacea: Isopoda:
Trichoniscidae) George A. Schultz
A new species of deep-sea isopod, Storthyngura myriamae, from the Walvis Ridge off South
Africa Robert Y. George and Noel A. Hinton
Phytoplankton distribution along the eastern coast of the USA IV. Shelf waters between Cape
Lookout, North Carolina, and Cape Canaveral, Florida Harold G. Marshall

Nomenclatural status of the foraminiferal genus Cubanella Saidova, 1981
Richard W. Huddleston and Drew Haman
Some new and old species of the primnoid genus Callogorgia Gray, with a revalidation of the

related genus Fanellia Gray (Coelenterata: Anthozoa) Frederick M. Bayer
Freshwater triclads (Turbellaria) of North America. XIII. Phagocata hamptonae, new species,
from Nevada Roman Kenk

Classificatory revisions in gammaridean Amphipoda (Crustacea), Part 2
J. Laurens Barnard and Gordon S. Karaman

Hemiodopsis ocellata, a new hemiodontid characoid fish (Pisces: Characoidea) from western

Surinam Richard P. Vari
Evidence of ontogenetic setal changes in Heteromastus filiformis (Polychaeta: Capitellidae)
Thomas J. Fredette
A new species of Ophiogomphus (Insecta: Odonata: Gomphidae) from the western Highland
Rim in Tennessee Jerry A. Louton
Two new species of Onuphis (Onuphidae: Polychaeta) from Uruguay Kristian Fauchald
Remarks on the Stylasterine fauna of the West Indies, with a description of Stylaster antillarum,

a new species from the Lesser Antilles (Cnidaria: Hydrozoa: Stylasterina)
Helmut Zibrowius and Stephen D. Cairns

48

58

67
81

89

93

99

114

116

161

167

188

194

198
203

210

Wag te Fag |
go -

324X)

(ISSN 0006
Number 2

t 1982

11 Augus

We Gee
SS SIBR

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i

THE BIOLOGICAL SOCIETY OF WASHINGTON

1981-1982
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PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 223-231

ATROPHECAECUM LOBACETABULARE, N. SP.
(DIGENEA: CRYPTOGONIMIDAE: ACANTHOSTOMINAE)
WITH DISCUSSION OF THE GENERIC STATUS OF
PARACANTHOSTOMUM FISCHTHAL AND KUNTZ, 1965,
AND ATEUCHOCEPHALA COIL AND KUNTZ, 1960

Daniel R. Brooks and Janine N. Caira

Abstract.—A new species of Atrophecaecum is described from various sea
snakes collected in northern Australia and Malaysia. The new species most
closely resembles A. burminis, A. simhai, and Paracanthostomum cerberi in
having one atrophied cecum, preovarian vitellaria, vitelline follicles not extending
anteriorly to the posterior margin of the seminal vesicle, eggs attaining lengths
greater than 30 um, and a short prepharynx. It is unique among acanthostomes
in possessing a flap of tissue on the posterior margin of the acetabulum, giving
that organ a bipartite appearance. The new species further resembles A. burminis
and-4.. simhai by possessing oral spines, and resembles P. cerberi in having a
subterminal mouth. Inclusion of Paracanthostomum cerberi and Ateuchocephala
marina in Atrophecaecum provides the most parsimonious hypothesis of phylo-
genetic relationships among the taxa involved, and produces a more efficient and
stable classification.

Brooks (1980) recently revised the digenean subfamily Acanthostominae Poche
based on a phylogenetic analysis. In that monograph a new species of Atrophe-
caecum Bhalerao was discussed in terms of its effect on the classification of
acanthostomes and two other cryptogonimid genera, Paracanthostomum Fisch-
thal and Kuntz, and Ateuchocephala Coil and Kuntz. Specifically, the possibility
was raised that Ateuchocephala and Paracanthostomum could not be recognized
as natural (monophyletic) taxa unless Atrophecaecum was somehow subdivided.
This report describes the new species of Atrophecaecum and, based on phylo-
genetic analysis, shows that the most useful classification for the taxa involved
is one including Ateuchocephala and Paracanthostomum in the genus Atrophe-
caecum.

Specimens upon which the description of the new species is based were col-
lected from sea-snakes quick-frozen shortly after capture and kept frozen until
returned to the laboratory, where they were thawed and examined. Most were
partially thawed, eviscerated, and the viscera refrozen and shipped to the Uni-
versity of Alberta. Collected helminths were fixed with cold AFA, stained with
Harris’ hematoxylin and mounted in Permount (Fisher). Measurements are in
micra unless otherwise stated, with mean values (x) and number of observations
(n) for some traits; figures were drawn with the aid of a drawing tube.

Atrophecaecum lobacetabulare, new species
Figs. 14

Description (based on 34 specimens).—Body elongate, 4.32—12.30 mm long
(n = 27) by 0.24-0.75 mm wide (n = 27); widest point variable in hindbody; ratio

224 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-4. Atrophecaecum lobacetabulare: 1, Ventral view of holotype; 2, Close-up of posterior
end of paratype, showing position of seminal receptacle, which is not apparent in holotype; 3, Close-
up of ootype region; 4, Dorsal view of acetabular region of holotype, showing terminal genitalia.
Abbreviations: HD = hermaphroditic duct; LC = Laurer’s canal; MG = Mehlis’ gland; O = ovary;
SR = seminal receptacle; SV = seminal vesicle; U = uterus; VR = vitelline reservoir; VD = vitel-
line duct.

of body width to length 1:9.1-33.8 (x = 1:18.1,n = 27). Tegument spinose; extent
of spination not determined due to fixation methods. Eyespot pigment lacking.
Oral sucker cup-shaped, terminal with subterminal mouth, 154—297 (n = 24) long
by 143-231 (n = 24) wide, surrounded by single uninterrupted row of 24—26
spines; spines 22-38 (n = 20) long by 12-29 (n = 20) wide. Acetabulum 105-231
long (n = 23) by 95-209 wide (n = 23), with posterior lobe giving bipartite ap-
pearance; lobe not apparent in many specimens. Forebody 6.4—-14.1% of total
body length. Ratio of oral sucker width to acetabular width 1:0.64—1.07 (« =
1:0.84, n = 20). Prepharynx 60-242 long (n = 21), thin-walled. Pharynx barrel-
shaped, 108-148 long (n = 22) by 99-214 wide (n = 22). Ratio of oral sucker
width to pharyngeal width 1:0.50-0.89 (x = 1:0.71, n = 16). Ratio of acetabular
width to pharyngeal width 1:0.59-1.15 (x = 1:0.87, n = 16). Esophagus usually
extremely short, not measured. Cecal bifurcation less than 5% of total body length
pre-acetabular; ceca lined with epithelium, opening separately and laterally near
posterior end of body; one cecum atrophied.
Testes spherical to subspherical, smooth, tandem, not contiguous; anterior
testis 209-345 long by 132-208 wide, posterior testis 220-396 long by 132-352
wide; post-testicular space 3—11% of total body length (x = 7%, n = 26). Seminal
vesicle consisting of posterior saccate part and anterior sinus part, sinuous, me-
dian, intercecal, extending 4.6—11.7 times acetabular length postacetabular (x =
9.1,n = 18). Prostatic duct surrounded by few prostatic cells free in parenchyma.
Preacetabular pit without spines, with transverse aperture 80-83 wide, surround-

VOLUME 95, NUMBER 2 UES)

ed by gland cells; gonotyl lacking. Genital pore immediately preacetabular, open-
ing immediately posterior to preacetabular pit. Postacetabular pit lacking. —

Ovary approximately one ovarian diameter pretesticular, not contiguous with
anterior testis, spherical to subspherical, 120-264 long by 99-242 wide. Seminal
receptacle posterodorsal to ovary, 132-506 long by 77-198 wide. Ootype region
as in Fig. 3. Uterus wound in ascending loops intercecal between ovary and
acetabulum; loops occupying 63-75% of total body length (x = 70%, n = 23);
joining hermaphroditic duct posterior to acetabulum. Vitellaria follicular; follicles
in 2 longitudinal rows dorsal and lateral to ceca, extending from immediately
preovarian to 8—50% of total body length postacetabular (x = 27%, n = 24), ter-
minating at uneven levels. Left vitellarium follicles extending anteriorly to 8-46%
of total body length postacetabular, right vitellarium follicles extending anteriorly
to 9-50% of total body length postacetabular. Follicles not reaching anteriorly to
posterior margin of seminal vesicle, 14—33 long by 12—29 wide. Eggs 25-35 (x =
28, n = 34) long by 13-20 (x = 16.5, n = 34) wide. Excretory vesicle Y-shaped;
bifurcation posterodorsal to acetabulum; pore terminal with muscular sphincter
surrounded by gland cells. |

Hosts.—Disteira major (Shaw), Hydrophis caerulescens (Shaw), H. spiralis
(Shaw), Enhydrina schistosa (Daudin) (type), Lapemis hardwicki Gray (Ophidia:
Hydrophiidae: Hydrophiinae: Hydrophiini).

Site of infection.—Small intestine.

Localities.—Penang, Malaysia (type) (E. schistosa, H. spiralis, L. hardwicki);
Western Gulf of Carpenteria, Australia (D. major, H. caerulescens).

Holotype.—USNM Helm. Coll. No. 77161. Paratypes.—USNM Helm. Coll.
No. 77162, 77163, 77164.

Etymology.—The species is named for its lobate acetabulum, a feature unique
among acanthostomes.

Atrophecaecum lobacetabulare most closely resembles the species A. bur-
minis (Bhalerao, 1926) Khalil, 1963, and A. simhai Khalil, 1963, and Para-
canthostomum cerberi Fischthal and Kuntz, 1965, by having one atrophied cecum,
preovarian vitellaria, vitelline follicles not extending anteriorly to the pos-
terior margin of the seminal vesicle, eggs attaining lengths greater than 30 um,
and a very short prepharynx. The new species is unique among known acan-
thostomes by virtue of its possessing a lobate acetabulum. It resembles A. bur-
minis and A. simhai by having oral spines, although they are relatively much
smaller than those of most other acanthostomes. It resembles P. cerberi by pos-
sessing a subterminal mouth with terminal oral sucker. Thus, A. lobacetabulare
occupies a systematic position intermediate between the taxa listed above.

Discussion

Brooks (1980) alluded to possible taxonomic problems arising from the discov-
ery of A. lobacetabulare. The nature of the problem, and of its solution, may be
seen best by examining the relative positions of A. lobacetabulare, P. cerberi,
and Ateuchocephala marina in a cladogram depicting the phylogenetic relation-
ships of Atrophecaecum spp. (Fig. 5). Synapomorphies (shared special traits)
linking the new species and the other two taxa above to Atrophecaecum are listed
on the cladogram (simplified from that of Brooks 1980).

Figure 5 clearly shows that A. lobacetabulare, P. cerberi, and Ateuchocephala

226 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ce (o)
ee? e&® >
yO .\ \\ CG C
so? SP oe Ron LO AS <S BS) ee ot
8 OXF of So © SS) x J
no prepharynx |
no esophagus short seminal vesicie
secondary vitelline clusters
lobate aground testes
acetabulum no oral spines (2° loss)
mouth subterminal
one cec :
eave uterine loops occupying more than 50% TBL

eggs longer than 30um
one cecum atrophied
short prepharynx

vitellaria terminating preovarially

Fig.5. Cladogram, partly modified from that of Brooks (1980), depicting phylogenetic relationships
of Atrophecaecum spp. plus Paracanthostomum cerberi and Ateuchocephala marina. Modifications
comprise deletion of some characters from the cladogram for the sake of clarity in viewing, and
addition of A. lobacetabulare and P. cerberi and A. marina.

marina form a monophyletic group most closely related to Atrophecaecum bur-
minis and A. simhai, hence our choice of comparisons above. This arrangement
would require that Paracanthostomum and Ateuchocephala be considered junior
synonyms of Atrophecaecum if A. lobacetabulare and all other members of the
genus are to be retained in a single genus. Despite the perceived morphological
distinctness of oral structures possessed by P. cerberi and Ateuchocephala ma-
rina, they represent only secondary loss of traits (oral spines) already reduced in
size in their closest relative and a tilting of the mouth orientation. Paracantho-
stomum and Ateuchocephala are therefore characterized in part by the absence
of traits. Conversely, Atrophecaecum would not exist as a natural taxon if P.
cerberi and Ateuchocephala marina are excluded from it.

Three types of objections commonly raised concerning cladistic classification
include: (1) they involve loss of information about ‘‘gaps’’ or degrees of difference
between taxa (anagenetic information), (2) they produce confusing and unusable
diagnoses and keys, and (3) any attempts to preserve cladistic information pro-
duce long unwieldy classifications. All of these objections have been treated
empirically in papers published by various authors primarily in Systematic Zo-
ology during the past ten years. This study provides an opportunity to demon-
Strate the efficacy of phylogenetic classification using a set of real taxa.

Loss of information.—Consider the three genera Atrophecaecum, Paracan-
thostomum, and Ateuchocephala in the classification presented by Yamaguti
(1971). Paracanthostomum and Atrophecaecum (as a subgenus of Acanthosto-
mum) belong in the Acanthostominae and Ateuchocephala comprises the sole

VOLUME 95, NUMBER 2 My]

member of the Ateuchocephalinae. Such a classification does not represent any
information concerning any members of Atrophecaecum which exhibit traits in-
termediate between the other taxa, nor does it recognize any of the similarities
between Ateuchocephala and Paracanthostomum. Yamaguti’s classification pro-
posed a set of relationships opposite to the phylogenetic relationships supported
by known data; Ateuchocephala was considered the sister-lineage of Paracan-
thostomum plus all other acanthostomes, and Paracanthostomum was considered
the sister-lineage of all armed acanthostomes, including Atrophecaecum. Even
if Paracanthostomum were considered a member of the Ateuchocephalinae, the
classification would suggest that Atrophecaecum excluding Paracanthostomum
and Ateuchocephala comprises a monophyletic group. Such a notion is not sup-
ported by any known data. In such cases, because Atrophecaecum lobacetabu-
lare possesses traits intermediate between Atrophecaecum and Paracanthosto-
mum plus Ateuchocephala, the “‘gap’’ between the taxa no longer exists, and
existed originally only as an artifact of sampling error. Insofar as gaps represent
either large numbers of character differences, which may be an artifact of data
type or of sampling, or large-scale character differences, which may also be an
artifact of data type, information concerning such occurrences should not be a
basis for classification. Such occurrences are represented in every cladogram but
do not alter the branching pattern or classification produced. A cladistic classi-
fication does not indicate directly such gaps, but the diagnoses for which the
taxon names stand do.

Confusion in diagnoses and keys.—It has been suggested (Mayr 1969; Sneath
and Sokal 1973) that cladistic classifications are not very stable. One aspect of
that instability would be wholesale changes in diagnoses and keys every time a
new taxon is included in the classification. It is true that the inclusion of the taxa
in question within Atrophecaecum requires changes in some diagnoses and keys.
However, we think the changes are few and minor, especially when compared
with the number of unnecessary redundancies in diagnoses required if Paracan-
thostomum and Ateuchocephalus were maintained separately from the other
acanthostomes.

Inclusion of Paracanthostomum and Ateuchocephalus in the Acanthostominae
requires the following changes (in italics) in the subfamilial diagnosis presented
by Brooks (1980):

Acanthostominae Poche, 1926

Diagnosis (emended from that of Brooks 1980).—Body elongate, eyespotted
or not. Oral sucker terminal, with terminal or subterminal mouth, surrounded by
uninterrupted single row of spines or lacking spines. Prepharynx and esophagus
present and variable in length or lacking. Ceca extending to near posterior end
of body. Acetabulum median, embedded in parenchyma or enclosed in body fold.
Preacetabular pit present. Postacetabular pit present or lacking. Testes 2, interce-
cal, tandem or oblique, in hindbody. Seminal vesicle present. Cirrus sac lacking.
Gonotyl present or lacking. Genital pore immediately preacetabular, not in pre-
acetabular pit. Ovary spherical or subspherical, pretesticular in hindbody. Sem-
inal receptacle and Laurer’s canal present. Vitellaria follicular, in lateral fields in
hindbody. Uterus postacetabular, usually not extending postovarially. Eggs em-

228 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

bryonated, not filamented. Excretory vesicle Y- or V-shaped, with post- or pre-
acetabular bifurcation. Parasites of estuarine and freshwater fishes and reptilians.
Pantropical. Type-genus: Acanthostomum Looss, 1899.

Inclusion of Paracanthostomum and Ateuchocephala in Atrophecaecum re-
quires the following emendations to the generic diagnosis presented by Brooks
(1980):

Atrophecaecum Bhalerao, 1940

Ateuchocephala Coil and Kuntz, 1960:145—150.
Paracanthstomum Fischthal and Kuntz, 1965:124—136.

Diagnosis.—Acanthostominae. Tegumental spines uniform in size. Oral sucker
armed with single uninterrupted row of spines or lacking spines. Mouth terminal
or subterminal. Prepharynx and esophagus variable in length or one lacking.
Ceca opening separately and laterally; ceca not atrophied, one cecum atrophied
or one cecum lacking. Gonotyl lacking. Seminal vesicle coiled. Vitelline follicles
terminating preovarially; secondary cluster of follicles surrounding testes may be
present. Seminal receptacle posterodorsal to ovary. Uterine loops preovarian.
Excretory vesicle Y-shaped, with pre- or postacetabular bifurcation. Parasites of
crocodilians and ophidians. India, Pakistan, Burma, Malaysia, Australia. Type-
species: Atrophecaecum burminis (Bhalerao, 1926), Bhalerao, 1940.

The above additions to the diagnoses of Acanthostominae and of Atrophecae-
cum eliminate the need for separate diagnoses of Paracanthostomum, Ateucho-
cephala and the Ateuchocephalinae. Additionally, the phylogenetic relationships
of the taxa involved are represented in the classification and the new classification
differs little from the previous classification of the acanthostomes with the ex-
ception of the addition of three new species. Thus, the cladistic system is de-
monstrably more stable and less ambiguous than any other. The above additions
do not require any changes in the generic key presented by Brooks (1980) but do
require a new key to the species of Atrophecaecum, as follows:

la. Prepharynx less than 3 times longer than pharynx, one cecum partially

or completely atrophied, parasites of ophidians ..................... 3

1b. Prepharynx at least 3 times longer than pharynx, ceca not atrophied,
parasites: of icrocodilians 277 ae oe ee Ce ee eee yy
2a. Oral:spines “8-19 ins num betes ernerceemoe Ae esos wee see slusarskit
2bz (Oral spimes*22: 1 number we we ee rs ee ee eee ae indicum
3a; One Cecum lacking” oY anet une retro cert ere or Ie a eee tr enn ee 4
3b.’ Both: Ceca present 0 eset eee se ee eee oe ee ee 5

4a. Oral spines 20-22 in number, vitellaria confluent preovarially
RSS TN ci A aA I CRE BS A I Mates sR gr OL ano. ce welt Hed proctophorum

4b. Oral spines 24—28 in number, vitellaria not confluent preovarially
CeO POEM k PONIES AN 1S Bra AI ciel oh rai la ood 0, 9 Souci asymmetricum

Sa. Oral spines 23 in number, vitelline follicles extending anteriorly to pos-
terior margin of seminal vesicle’ 9... -22-72-- 20-2 s0- ee 4 pakistanense

5b. Oral spines 24—29 in number or lacking, vitelline follicles not extending
anteriorly to posterior margin of seminal vesicle, eggs reaching more
than-30" am in'length ss: Seer ee ee cre oe ee te cece pect 6

VOLUME 95, NUMBER 2 229

6a. Cecal bifurcation more than 10% of total body length preacetabular,

uterine loops occupying less than 50% of total body length ........ simhai
6b. Cecal bifurcation less than 5% of total body length preacetabular, uterine
loops occupying more than 50% of total body length ................. 7
Fal, IMIOWICI USI aie ener ine e . | is burminis
FP MOMUMESMOLETIMIMAL, Js cc... occ gate ee ee ke ee Hs Seis bs sce ea eas 8
8a. Oral spines present, acetabulum bipartite ................ lobacetabulare
8b. Oral spines lacking, acetabulum unipartite ............ 0.0.0. e cee eee 9
9a. Esophagus present, prepharynx lacking, secondary vitelline cluster sur-
hoanmems testes, Seminal) vesicle: Short’) (228) ote ee ee marinum
9b. Esophagus lacking, prepharynx short, secondary vitelline cluster lack-
imenseminaly vesicle elongate .):c00. on. s oe 5 oes we ieee Pees 2 cerberi

Coil and Kuntz (1960) reported lateral ani for Ateuchocephala marina, but
Yamaguti (1971) reported no ani after examining a paratype. We examined the
specimens in question and found lateral ani and slight cecal atrophy. Yamaguti
further reported a fairly long prepharynx in Paracanthostomum cerberi, but we
found the prepharynx of the holotype and of specimens reported by Brooks and
Palmieri (1981) to be as short as or shorter than the pharynx.

A number of species of Ateuchocephala have been described in addition to A.
marina, but their validity and identities remain in question pending acquisition
of material for study. Therefore, we make only the following new combinations:
Paracanthostomum cerberi Fischthal and Kuntz, 1965 becomes Atrophecaecum
cerberi (Fischthal and Kuntz, 1965), n. comb., and Ateuchocephala marina Coil
and Kuntz, 1960 becomes Atrophecaecum marinum (Coil and Kuntz, 1960), n.
comb.; Ateuchocephalinae Yamaguti, 1971 becomes a junior subjective synonym
of Acanthostominae Poche, 1926.

Inclusion of all cladistic information in classifications produces long unwieldy
classifications.—Do attempts to preserve all the information in a cladogram nec-
essarily produce long unwieldy classifications? In order to examine this question,
we formulated a complete classification of the acanthostomes according to the
convention of phyletic sequencing (see Wiley 1979) in which every taxon is con-
sidered the sister-group of all taxa of equivalent position below it. That classifi-
cation:

Subfamily Acanthostominae
Genus Timoniella
Subgenus 1
species praeterita
species imbutiformis
Subgenus 2
species incognita
species scyphocephala
species unami
species loossi
Genus Gymnatrema
species gymnarchi
species pambanense
Genus Proctocaecum
Subgenus 1
Infrasubgenus 1

230 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

species coronarium
species vicinum
Infrasubgenus 2
species productum
species gonotyl
Subgenus 2
Infrasubgenus 1
species nicolli
Infrasubgenus 2
species atae sedis mutabilis
species elongatum sedis mutabilis
species crocodili sedis mutabilis
Gerwus Caimanicola
species caballeroi
species pavidus
species marajoara
species brauni
Genus Acanthostomum
Subgenus 1
species absconditum
species knobus
species niloticum
species spiniceps
Subgenus 2
Infrasubgenus 1
species gnerii
species minimum
species astorquii
Infrasubgenus 2
species americanum
species megacetabulum
Genus Atrophecaecum
Subgenus 1
species indicum
species slusarskii
Subgenus 2
species pakistanense
Subgenus 3
species proctophorum
species asymmetricum
Subgenus 4
species simhai
species burminis
species lobacetabulare
species cerberi
species marinum

This classification is only 16 lines longer than a classification listing only genera
and their included species, and only 6 lines longer than a classification using
subgeneric designations. Thus, a fully-resolved cladistic classification need not be
substantially longer than a less informative syncretistic scheme.

This report demonstrates the utility of a cladistic classification scheme as a
general reference system for helminth systematics. Theoretical claims in support
of the stability, consistency, predictivity, information content, and applicability
of Hennigian systematics are upheld in this practical demonstration. We strongly

VOLUME 95, NUMBER 2 Vip)

urge helminthologists to investigate the use of such an approach in classifying
other taxa.

Acknowledgments

We express appreciation to Dr. John C. Holmes, University of Alberta, for pro-
viding the specimens of Atrophecaecum lobacetabulare. Funds for this study were
provided through a grant to DRB from the Natural, Applied and Health Sciences
Grant Committee of the University of British Columbia.

Literature Cited

Brooks, D. R. 1980. Revision of the Acanthostominae Poche, 1926 (Digenea: Cryptogonimidae).—

Zoological Journal of the Linnean Society 70:313-382.

, and J. R. Palmieri. 1981. Six platyhelminths from Malaysian reptiles including Paradisto-

moidella cerberi n. g., n. sp. (Digenea: Dicrocoeliidae)—Journal of Helminthology 55:39-43.

Coil, W. H., and R. E. Kuntz. 1960. Three new genera of trematodes from Pacific sea serpents.—
Proceedings of the Helminthological Society of Washington 27: 145-150.

Fischthal, J. H., and R. E. Kuntz. 1965. Digenetic trematodes of amphibians and reptiles from North
Borneo (Malaysia).—Proceedings of the Helminthological Society of Washington 32:124—-136.

Mayr, E. 1969. Principles of systematic zoology.—McGraw-Hill, New York.

Sneath, P. H. A., and R. R. Sokal. 1973. Numerical taxonomy.—W. H. Freeman and Co., San
Francisco.

Wiley, E. O. 1979. An annotated Linnean hierarchy, with comments on natural taxa and competing
systems.—Systematic Zoology 28:308-337.

Yamaguti, S. 1971. Synopsis of digenetic trematodes of vertebrates.—Keigaku Publ. Co., Tokyo.

(DRB) Department of Zoology, University of British Columbia, 2075 Wesbrook
Mall, Vancouver, B.C. V6T 2A9, Canada; (JNC) Harold W. Manter Laboratory,
Division of Parasitology, University of Nebraska State Museum, W-529 Nebraska
Hall, Lincoln, Nebraska 68588-0514.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 232-237

A PARTIAL REVISION OF THE GENUS NOTOMASTUS
(POLYCHAETA: CAPITELLIDAE) WITH A DESCRIPTION OF
A NEW SPECIES FROM THE GULF OF MEXICO

R. Michael Ewing

Abstract.—An emended diagnosis for the genus Notomastus (Polychaeta: Cap-
itellidae) is proposed with two closely related genera, Rashgua Wesenberg-Lund
and Paraleiocapitella Thomassin, considered as junior synonyms. A new species
of Notomastus from the Gulf of Mexico, N. daueri, is described.

Recent benthic ecological studies in the northern Gulf of Mexico revealed that
an undescribed capitellid polychaete was a prominent component of the infauna
of relatively shallow silty sand habitats. Description of this species necessitated
expansion of the generic diagnosis of Notomastus Sars, 1851. Close examination
of the literature indicated that two closely related genera, Rashgua Wesenberg-
Lund, 1949 and Paraleiocapitella Thomassin, 1970 are sufficiently similar to be
considered as synonyms of Notomastus.

The holotype and one set of paratypes of the new species are deposited in the
National Museum of Natural History (USNM), Smithsonian Institution, Wash-
ington, D.C. An additional paratype is deposited in Allan Hancock Foundation
(AHF), University of Southern California, Los Angeles, California.

Family Capitellidae Grube, 1862
Genus Notomastus Sars, 1851, emended

Rashgua Wesenberg-Lund, 1949:336.
Paraleiocapitella Thomassin, 1970:86.

Type-species.—Notomastus latericeus Sars, 1851:199. |

Diagnosis.—TYhorax with an achaetous peristomium and 11 setigerous seg-
ments; epithelium wholly or partly smooth, tessellated, or areolated. Prostomium
conical or triangular, with or without palpode; eyespots present or lacking. First
setiger with or without capillary setae in neuropodia; thereafter thoracic segments
with capillary setae only in both rami or with hooks in neuropodia of last 1-3
setigers. Nephridial apertures absent, limited to either thorax or abdomen, or
present on a few segments in both regions. Abdominal neuropodia with hooded
hooks only; notopodia with hooks only, present throughout the abdomen, re-
stricted to anterior segments, or completely absent. Branchiae present or lacking;
if present, as nonretractile, simple expansions or prolongations of noto- and/or
neuropodia or as eversible branched tufts from notopodial ridges. Pygidium with
or without appendages.

Discussion

Remarks on Rashgua Wesenberg-Lund, 1949: Wesenberg-Lund (1949:336)
erected the genus Rashgua, characterized as having a thorax consisting of an

VOLUME 95, NUMBER 2 238

achaetous peristomium and 11 segments with capillary setae only in both noto-
and neuropodia and an abdomen with hooded hooks only. Wesenberg-Lund pre-
sumably distinguished Rashgua from the closely related genus Notomastus on
the basis that the former genus had abdominal notopodia “‘perfectly devoid of
hooks’’ (1949:337) while the latter had (by original definition) hooks in both rami
of the abdomen. However, Hartman (1947:415) described a species of Notomas-
tus, N. lobatus, which agrees with Rashgua in lacking abdominal notosetae.
Although Hartman (1947:416) did not emend the generic diagnosis of Notomastus
to accommodate N. lobatus, her observation that ‘‘notopodial tori are believed
to be absent”’ was apparently overlooked by Wesenberg-Lund (1949).

The original description of the genotype of Rashgua, R. rubrocincta, was based
on anterior fragments only. This species agrees with N. lobatus in most characters
but, based strictly on the literature account of R. rubrocincta, the 2 appear to
differ slightly in the configuration of abdominal tori; these 2 species are the only
known capitellids completely lacking abdominal notosetae.

A new species of Notomastus described in this paper has notopodial hooks in
anterior abdominal segments only. This structure is intermediate between the
conditions of no abdominal notosetae (Rashgua and Notomastus lobatus) and
notosetae throughout the abdomen (all other known Notomastus). The presence
or absence (complete or partial) of notosetae in the abdomen is now recognized
as a variable character in the genus Notomastus. Therefore, the genus Rashgua
is herein designated a junior synonym of Notomastus Sars. Notomastus rubro-
cinctus (Wesenberg-Lund, 1949) is considered a new combination.

Remarks on Paraleiocapitella Thomassin, 1970: Thomassin (1970:86) erected
the genus Paraleiocapitella which is characterized as having a thorax with an
achaetous peristomium and 11 setigerous segments; an incomplete first setiger
(i.e. notopodia only) is followed by 9 setigers with capillary setae only in both
rami and a last thoracic segment with capillary setae only in the notopodia and
hooded hooks only in the neuropodia; abdominal segments are provided with
hooks only in both noto- and neuropodia.

Paraleiocapitella was presumably easily distinguished from the nearly identical
genus Notomastus based on the presence of hooks in thoracic neuropodia in the
former genus and by original description, capillary setae only in the thorax of
Notomastus. However, prior to 1970, descriptions of at least 2 species of No-
tomastus with thoracic neuropodial hooks had been published although the ge-
neric description was not technically revised. Notomastus precocis Hartman,
1960, has hooks only in the last 3 thoracic neuropodia. Hartman (1965) also
described Notomastus teres which is characterized as having hooded hooks only
in the last 2 neuropodia. Day (1973), in agreement with Hartman’s expanded
diagnosis of Notomastus but seemingly unaware of the genus Paraleiocapitella,
described Notomastus americanus which has a thoracic setal formula identical
to that of the genotype of Paraleiocapitella, P. mossambica.

In the present study, numerous immature specimens representing 3 additional
species of Notomastus, N. hemipodus Hartman, 1947, N. lobatus Hartman, 1947,
and N. daueri, n. sp., were found with hooded hooks only or mixed fascicles of
capillary setae and hooks in as many as 5 posterior thoracic neuropodia and rarely
in 1-2 notopodia. With very few exceptions, a distinct pattern of setal develop-
ment toward the adult arrangement (i.e. thoracic setal formula) was recognized

234 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

in each of these species. Similar conditions were also noted in juveniles of several
other capitellid genera. The subject of setal development will be discussed in
greater detail in a forthcoming paper. Tentatively outlined, the replacement of
hooded hooks by capillary setae in the thoracic neuropodia of Notomastus is as
follows: 1) juveniles are provided with hooks only in neuropodia of several seg-
ments in the posterior one-half of the thorax, 2) as the worm grows, hooks are
apparently lost (shed, broken or resorbed?) from the ventral position in the setal
fascicle and replaced by capillary setae emerging from the superior position; the
replacement of setae progresses until all hooks in a ‘‘changing’’ fascicle are re-
placed by capillary setae and the process continues in a relatively predictable
manner toward the posterior thoracic segments until the adult condition is
reached.

In summary, the presence and/or location of hooded hooks in thoracic neuro-
podia of both juveniles and adults is recognized as a highly variable character in
the genus Notomastus and perhaps many other capitellid genera; thus the genus
Paraleiocapitella Thomassin is herein designated a junior synonym of Notomas-
tus Sars. Notomastus mossambicus (Thomassin, 1970) will be considered a new
combination pending personal examination of type-material.

Notomastus daueri, new species
Fig. la—g

Material examined.—LOUISIANA: Gulf of Mexico: Approx. 29.3 km SSW
Grand Isle: 28°56'12”N, 90°04'07’W, 27.7 m, silty clay, holotype (LW&F, col.,
16 April 1980, USNM 71442), 2 paratypes (USNM 71443), 18 specimens; 8 spec-
imens, 21 Aug. 1980, same location; 2 specimens, 8 Sept. 1980, same location.
Approx. 33.9 km SSW Grand Isle: 28°53’06"N, 90°01'30”W, 33.5 m, clayey silt,
1 paratype (LW&F, col., 8 Sept. 1980, AHF Poly 1361). Approx. 37.0 km SSW
Grand Isle: 28°51'06’N, 90°04'21”W, 33.2 m, 1 specimen, 21 Aug. 1980. MISSIS-
SIPPI: Gulf of Mexico: IEC 732 MO Sta. 014, approx. 5.9 km S Ship Island pass,
30°10.32’N, 88°55.00’W, 10 m, muddy sand, 1 specimen (IEC, col., June 1980);
Mississippi Sound: sta. 043, approx. 3.6 km N Petit Bois Island, 30°14.48’N,
88°25.63’W, 5.6 m, muddy sand, 1 specimen (Vittor & Assoc., col., 22 Oct. 1980).

Description.—Length of largest complete specimen (holotype) approximately
65 mm, width 1.1 mm, 234 setigerous segments. Lengths of 7 additional complete
specimens ranged from 40 to 52 mm, widths 0.3 to 0.9 mm, with up to 197 setigers.

Color light tan to brown in alcohol. Thorax slightly inflated through setigers 4—
5, surface epithelium faintly areolated; following thoracic segments nearly smooth
except for ventral biannulation. Abdominal epithelium smooth throughout.

Prostomium short, broadly triangular in dorsal view, with 2 inconspicuous nu-
chal slits near posterior border; eyespots absent. Achaetous peristomium slightly
wider than long, approximately 114 to 2 times as long as following segment;
eversible pharynx bulbous, coarsely papillated on proximal two-thirds, smooth
distally. Thoracic setigers about 3 times as wide as long. Anteriormost notopodia
dorsolateral, well separated, but approach middorsally by setigers 6—7; neuropodia
ventrolateral in position throughout thorax. Anterior 10 setigers with 15-20 cap-
illary setae per fascicle in both rami; last thoracic segment (setiger 11) with cap-
illary setae only in notopodia and fascicles of 12-20 hooded hooks only in neu-
ropodia (Fig. la). Nephridial apertures (1 pair on each segment) located in

VOLUME 95, NUMBER 2 235

0.025mm

0.5mm d

Fig. 1. Notomastus daueri: a, Lateral view of anterior end showing thorax and first 2 abdominal
segments; b, Dorsolateral view of midabdominal segments; c-d, Lateral and dorsal views of posterior
abdominal segments; e, Dorsal view of pygidium; f—g, Lateral and frontal views of neuropodial hooded
hook from midabdomen.

segmental groove between each of last 4 thoracic segments. Lateral organs pres-
ent on all thoracic setigers as a minute pore between noto- and neuropodium;
appearing as a small ovoid structure just above neuropodial tori in abdominal
segments, present at least through midabdomen but becoming increasingly less
conspicuous moving posteriorly. Transition from thorax to abdomen marked by
setael change in notopodia from capillary setae to hooks, increase in number of
neurosetae per fascicle and slight broadening of segments.

Anterior abdominal segments of approximately same length as those of the
posterior thorax, gradually lengthening to the midabdominal region where they
are 114 to 2 times as long as wide; thereafter segments becoming increasingly
shorter.

Branchial lamellae first emerging from anterior abdominal segments (about se-
tiger 60 in holotype) as short posteriolateral swellings of the notopodia, increasing
in length posteriorly (Fig. 1b), developing into large triangular projections which
overlap the following segment in the posterior region (Figs. I1c—d).

Abdominal parapodia with multidentate hooded hooks only; hooks consisting

236 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

of a main fang surmounted by 3 teeth in a triangular arrangement and a second
row of 4—5 smaller denticles (Figs. le—f), similar in structure to those in the last
thoracic neuropodia. Anteriormost notopodia with 15-20 setae per fascicle on
short, slightly elevated tori; notosetae decreasing rapidly in number posteriorly,
disappearing completely 5-15 segments after the appearance of branchial lobes.
Neurosetal fascicles on well-defined glandular ridges at posterior margin of each
segment, extending from lateral to near midventral position; neurosetae increas-
ing in number posteriorly to as many as 40 hooks per fascicle by midabdomen,
then decreasing in number through posterior abdomen to 1-2 per fascicle in last
8-10 segments; noticeable reduction in size (and perhaps number) of denticles
above the main fang of hooks in far posterior region with some hooks appearing
unidentate under oil immersion in last few segments. Neuropodial tori of latter
segments enlarged at posterior margin as a shallow, somewhat cup-shaped pro-
cess (dorsal view) which may also be branchial in function.

Pygidium funnel-shaped with single digitate median cirrus (Fig. 1g).

Remarks.—Variations from the thoracic setal arrangement of adults were ob-
served in numerous small specimens (Guveniles?) of Notomastus daueri.

Notopodia of setigers 10 and 11 were rarely found with mixed fascicles of
capillary setae and hooks. Neuropodia of setigers 7-11 may have capillary setae
only, mixed setal fascicles, hooded hooks only, or combinations of these in a
given segment.

Notomastus daueri differs from most other species of the genus in having some
thoracic neuropodia with hooded hooks instead of capillary setae only. The 4
known species of Notomastus with hooks in one or more thoracic neuropodia,
N. precocis Hartman, 1960, N. teres Hartman, 1965, N. mossambicus (Thomas-
sin, 1970; new combination proposed in this paper), and N. americanus Day,
1973, all have notopodia only on setiger 1 whereas N. daueri has a complete first
setiger.

Notomastus daueri is known to occur in the northern Gulf of Mexico off Lou-
isiana and Mississippi in shallow subtidal muddy sands.

Etymology.—This species is named in honor of Dr. Daniel M. Dauer, my good
friend and former graduate advisor, in an attempt to express my sincere appre-
ciation for his unselfish guidance over the past several years.

Acknowledgments

Some of the specimens examined in this study were collected at stations selected,
sampled and processed under the supervision of John deMond of Louisiana Wild-
life and Fisheries under contract to LOOP, Inc. Special thanks are extended to
the Seafood Division (LW&F) and A. J. Heikamp (LOOP, Inc.) for permission
to use this material. Additional specimens were obtained from Interstate Elec-
tronics Corporation (IEC) under contract no. 68-01-4610 from the Environmental
Protection Agency.

Barry A. Vittor & Associates, Inc. provided the laboratory space and materials
used in this study and I sincerely thank the following employees for their con-
tributions: Caroline Coker, Michael Milligan, Donna Valentine and Jewel Walt-
man assisted in assembling the collection. Jerry Gathof, Gary Goeke, and Paul
Wolf commented on the manuscript. Figures were prepared for press by Ben
Jordan. Several drafts of the paper were typed by Dottie Cotton.

VOLUME 95, NUMBER 2 743) ])

I also thank Susan Williams of Allan Hancock Foundation for arranging loans
of type-material.

My studies of capitellid polychaetes have benefitted greatly from discussions
with Dr. Kristian Fauchald of the Smithsonian Institution. I am especially ap-
preciative of his many helpful suggestions and of his constructive criticism of
this manuscript.

Literature Cited

Day, J. H. 1973. New Polychaeta from Beaufort, with a key to all species recorded from North
Carolina.—National Marine Fisheries Service Circular 375:1—140.

Grube, A. E. 1862. Noch ein Wort tiber die Capitellen und ihre Stellung im Systeme der Anneliden.—
Archiv ftir Naturgeschichte (Berlin) 28:366-378.

Hartman, O. 1947. Polychaetous annelids. Pt. 7. Capitellidae.—Allan Hancock Pacific Expeditions

10:391-481.

. 1960. Systematic account of some marine invertebrate animals from the deep basins off

Southern California.—Allan Hancock Pacific Expeditions 22:69-216.

. 1965. Deep-water benthic polychaetous annelids off New England to Bermuda and other

North Atlantic areas.—Allan Hancock Foundation Publications Occasional Paper 28: 1-378.

Sars, M. 1851. Beretning om en i Sommeren 1849 foretagen zoologisk Reise i Loften og Fimmarken.—
Nyt Magazin for Naturvidenskaberne Christiania 6:121-211.

Thomassin, B. 1970. Contribution a l’étude des polychetes de la region de Tuléar (SW de Mada-
gascar). Sur les Capitellidae des sables corralliens.—Recueil des Travaux de la Station Marine
d’Endomme, Supplément 10:71-101.

Wesenberg-Lund, E. 1949. Polychaetes of the Iranian Gulf.—Scientific Investigations in Iran 4:247-—
400.

Barry A. Vittor & Associates, Inc., 8100 Cottage Hill Road, Mobile, Alabama
36609. Present address: Department of Biological Sciences, Old Dominion Uni-
versity, Norfolk, Virginia 23508.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 238-250

SOME SPECIES OF ONUPHIS (POLYCHAETA: ONUPHIDAE)
FROM THE ATLANTIC OCEAN

Kristian Fauchald

Abstract.—Five new species of Onuphis (Polychaeta: Onuphidae) are de-
scribed. Three, O. (Nothria) australatlantica, O. (N.) heterodentata, and O.
(N.) lithobiformis come from southern Atlantic waters; the other two, O. (On-
uphis) declivorum, and O. (O.) texana are from the east coast of the United
States.

Introduction

A review of material of onuphids deposited in the collections of the National
Museum of Natural History, Smithsonian Institution, has revealed the presence
of five previously undescribed species. Part of the material was previously treated
by Hartman (1967) in her study of Antarctic polychaetes; other specimens come
from benthic surveys along the east coast of the United States.

Generic definitions and terminology are those in Fauchald (1980).

Family Onuphidae Kinberg, 1866
Genus Onuphis Audouin and Milne Edwards, 1833
Subgenus Nothria Malmgren, 1866
Onuphis (Nothria) australatlantica, new species
Fig. 1, Table 1

Nothria cf. conchylega.—Hartman, 1967:91 (not M. Sars, 1835:61).

Material examined.—Antarctic Ocean, off South Shetland Islands, 3768—3816
m, Menzies-trawl, 2 August 1962, Eltanin Sta. 129 (17 paratypes, USNM 58296).
Antarctic Ocean, off South Shetland Islands, 3138-3239 m, Blake-trawl, 8 Sep-
tember 1963, Eltanin Sta. 722 (holotype, USNM 58195, one paratype, USNM
67488).

Description.—The holotype is an incomplete specimen with 29 setigers that is
23 mm long and 2.75 mm wide, with parapodia; it is a mature female with large
eggs in the body cavity. The body is dorsoventrally flattened with the parapodia
attached at the margins. All types are paie pink and lack color patterns. Eyes are
absent. The holotype is described in detail below; a summary of information for
all types is given in Table 1.

The prostomium (Fig. 1b) is rounded frontally and is wider than long. The
frontal palps are shorter than the length of the prostomium. The outer lateral
occipital antennae reach setiger 1, the inner lateral antennae reach setiger 4 and
the median antenna reaches setiger 7. The occipital ceratophores are short and
have up to 3 rings. Branchiae are absent.

The first 2 setigers (Fig. la) are enlarged. The first parapodia are directed
forward and laterally so that the morphologically anterior face of each parapodium
is medial. In the first pair of parapodia (Fig. 1li-j) the flattened, auricular presetal

VOLUME 95, NUMBER 2 7339)

Fig. 1. Onuphis (Nothria) australatlantica (holotype, USNM 58195): a, Anterior end, lateral view;
b, Anterior end, dorsal view; c, Pseudocompound hook, second setiger; d, Intrafascicular hook,
median setiger; e, Pectinate seta, median setiger; f, Pseudocompound hook, first setiger; g, Second
parapodium, anterior view; h, Second parapodium, posterior view; i, First parapodium, posterior
view; j, First parapodium, anterior view. All scales are 1 mm except where otherwise indicated.

240 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Summary statistics for Onuphis (Nothria) australatlantica.

Range Mean SD V N

Occipital antennae
Outer lateral reach # 1 invanant 19
Inner lateral reach # 4-8 5.89 1.08 1.16 18
Median reaches # 6-10 8.14 3) 1.52 14
# of rings 23 2.24 0.44 0.19 19
Ventral cirri cirriform to setiger # D invanant 19
Pseudocompound hooks to setiger # 2-3 2.05 0.23 0.05 19
Intrafascicular hooks from setiger # 9-11 9.95 0.71 0.50 19

lobe enfolds the dorsal side of the superiormost setae. The digitiform postsetal
lobe is directed posteriorly. The tapering dorsal cirrus is about twice as long as
the postsetal lobe. The ventral cirrus is similar to the postsetal lobe in size and
shape (the postsetal lobe projects posteriorly and is thus foreshortened in Fig.
li).

The second parapodia (Fig. 1g—h) are shorter than the first and directed later-
ally. The presetal lobe is less than half as wide as that of the first setiger, but is
still distinctly flattened and auricular. The postsetal lobe is about as long as that
of the first setiger, but is tapering from the base, rather than digitiform. The dorsal
cirrus resembles the one in the first setiger. The ventral cirrus is very short and
sharply tapering.

All other parapodia resemble each other; the presetal lobes are completely
reduced and the ventral cirri are replaced by glandular pads. The postsetal lobes
become increasingly threadlike towards the posterior end, but are distinct in all
setigers present; the dorsal cirri also become increasingly slender towards the
posterior end.

Pectinate setae, limbate setae, and anterior pseudocompound and intrafasci-
cular hooks are present. Compound spinigers, large hooks, and subacicular hooks
are absent. Pectinate setae (Fig. le) are present from the second setiger; each
seta is distally scoop-shaped and has about 15 teeth. About 12 pectinate setae are
present per parapodium in the anterior third of the body. The limbate setae have
extremely long shafts with a short, slightly geniculate limbation distally. The
pseudocompound hooks of the first setiger (Fig. 1f) are very thick, indistinctly
bidentate, and have short, blunt hoods. The hooks of the second setiger are less
than half as wide as those of the first setiger (Fig. 1c); the hinge-line is distinct
and each hook is clearly bidentate with a short, blunt hood. Pseudocompound
hooks are present in the first 2 setigers only. A pair of intrafascicular hooks is
present in each setiger from setiger 10. The intrafascicular hooks are oriented
parallel to the acicula and the limbate setae, and are about as long as the latter.
Each intrafascicular hook is bidentate distally and the head of the hook is tilted
so that both teeth are axial.

The maxillary formula (examined in the paratype from USNM 67488) is 1+1,
8+ 10, 3+0, 4+9 and 1+1; the unpaired maxilla HII has 2 poorly defined basal
teeth and a large distal fang.

The tubes are quadrangular in cross-section. The sides of the tubes that cor-

VOLUME 95, NUMBER 2 241

respond to the dorsal and ventral surface of the worm are somewhat more flat-
tened than the 2 lateral sides. The inner lining is tough; the outer surface is
covered with black, volcanic sand and other large particles that are not arranged
in a recognizable pattern.

Only one previously described species of Onuphis (Nothria) combines pro-
longed first parapodia with a lack of branchiae. Onuphis (N.) textor (Hartman
and Fauchald, 1971:78) was described from deep water in the Atlantic Ocean. It
is a very small species; a mature female with 15 setigers is about 2.5 mm long,
and has extremely long, slender first parapodia projecting well beyond the tip of
the prostomium. Onuphis (N.) australatlantica is a much larger and more massive
species; a mature female is 23 mm for 29 setigers, and the first parapodia, while
enlarged, do not project beyond the tip of the prostomium.

These specimens were originally identified as Nothria cf. conchylega by Hart-
man (1967); they differ clearly from Onuphis (Nothria) conchylega Sars, 1835, in
that they lack branchiae, present in the latter.

Etymology.—tThe specific name, australatlantica, refers to the localities in the
southern Atlantic Ocean from which material of the species was collected.

Distribution.—Onuphis (N.) australatlantica is known from two localities in
abyssal depths in the southern Atlantic Ocean.

Onuphis (Nothria) heterodentata, new species
Fig. 2, Table 2

Nothria ?iridescens.—Hartman, 1967:91 (in part, not Johnson, 1901:408).

Material examined.—South Atlantic Ocean, off Falkland Islands, 587-595 m,
rock dredge, 3 December 1962. Eltanin Sta. 338 (holotype, USNM 58299, 13
paratypes, USNM 67489).

Description.—The holotype is an incomplete specimen with 46 setigers, is 18
mm long and 1.5 mm wide, with parapodia. Other specimens are up to 24 mm
long for 55 setigers; none are complete. The anterior end of the specimens is
cylindrical; the dorsum is flattened and the ventrum is curved in median and
posterior setigers. The holotype (Fig. 2c) has lost the styles on both the inner
lateral and median occipital antennae. The outer lateral antennae reach setiger 2
and the ceratophores are distinctly longer than the styles. The inner lateral cer-
atophores, which are the longest, have up to 13 rings. In the paratypes, the inner
lateral antennae reach setiger 7 and the median antenna reaches setigers 4—5
(Table 2). The short frontal palps are ovate; eyes are absent.

Branchiae are present from setiger 2 in the holotype and from setigers | or 2 in
the paratypes. If branchiae are present from setiger 1, the first branchiae may be
missing from one side of the animal. Branchiae are present to the end of all
fragments; each is simple and straplike, and flattened, especially in posterior
setigers.

The first 2 pairs of parapodia are directed anteriorly. Ventral cirri are cirriform
in the first 5 setigers (Fig. 2f) and the postsetal lobes are digitiform in the first 11
setigers. The first 2 parapodia (Fig. 2a) have distinct contraction folds at the bases
of the dorsal cirri and rounded presetal and acicular lobes. The postsetal lobes
have thickened, pad-shaped bases. Median and posterior parapodia are barely
raised ridges with the setae emerging in a crescentic opening ventrally. The ven-

242 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

|]

h

Fig. 2. Onuphis (Nothria) heterodentata (holotype, USNM 58299): a, Second parapodium, an-
terior view; b, Pectinate seta, median setiger; c, Anterior end, dorsal view; d—e, Superior and inferior
pseudocompound hook, second parapodium; f, Anterior end, lateral view; g—h, Superior and inferior
pseudocompound hooks, third parapodium. All scales are 1 mm except where otherwise indicated.

tral glandular pads that replace the ventral cirri are very large and extend beyond
the tips of the subacicular hooks in most setigers.

Limbate and pectinate setae, pseudocompound and subacicular hooks are pres-
ent. Compound spinigers and large hooks are absent. Limbate setae are most
common in anterior parapodia except in the first 2 where only 1 or 2 are present.
One or 2 pectinate setae (Fig. 2b) are present in each parapodium; each pectinate
seta is slightly oblique distally, forming a 120° angle with the long axis of the seta,
and has 13-14 teeth; the 2 marginal teeth are not prolonged. Tridentate pseudo-
compound hooks (Fig. 2d—e, g—h) with short hoods are present in the first 3 or
4 parapodia. Only a single hook is present in setiger 4. The middle tooth projects
beyond the other teeth. The proximal tooth is broadly conical, the middle tooth
is curved, nearly beaked, and the distal tooth is thick and abruptly tapering

VOLUME 95, NUMBER 2 243

Table 2.—Summary of statistics of Onuphis (Nothria) heterodentata.

Range Mean SD V N

Occipital antennae

Outer lateral reach # D, invanant 12

Inner lateral reach # 5-10 WMG 2.14 4.57 6

Median reaches # 3-6 4.67 1,533 2.33 3

# of rings 12-13 12.20 0.42 0.18 10
Branchiae first present from setiger # 1-2 1.60 0.52 0.27 10
Ventral cirri cirriform to setiger # 5 invariant 14
Pseudocompound hooks present to

setiger # 4 invariant 14
Subacicular hooks present from

setiger # 12-14 12.50 0.65 0.42 12

distally. Two bidentate subacicular hooks are present in each parapodium from
setiger 12.

The maxillary formula (investigated in 2 paratypes) is 1+1, 9+9, 8—9+0,
7—8+9-—10 and 1+1.

Onuphis (N.) heterodentata resembles O. (N.) holobranchiata Marenzeller
(1879:132), O. (N.) iridescens (Johnson, 1901:408), O. (N.) opalina (Verrill,
1873:102), and O. (N.) lithobiformis (see below). All these species have simple
branchiae starting on setiger | and tridentate pseudocompound hooks exclusively.
Onuphis heterodentata differs from the other 4 species in that the median tooth
of the pseudocompound hooks is larger and projects further than the other 2
teeth; in all the other species the middle tooth is either of the same size as the
proximal tooth or intermediate in size between a long distal tooth and a short
proximal tooth. The relationships between the other 4 species are discussed be-
low.

Etymology.—The specific name refers to the different shapes of the teeth in
the pseudocompound hooks.

Distribution.—Onuphis (N.) heterodentata is known from a single locality in
slope-depths off the Falkland Islands.

Onuphis (Nothria) lithobiformis, new species
Fig. 3

Nothria ?iridescens.—Hartman, 1967:91 (in part, not Johnson, 1901:408).

Material examined.—South Atlantic Ocean, off Falkland Islands, from 53°08’S,
59°23'W to 53°07'S, 59°21'W, 578-567 m, Menzies-trawl, 3 December 1962, Eltanin
Sta. 340 (holotype, USNM 58300, 4 paratypes, USNM 67490). South Atlantic
Ocean, Straits of Magellan, from 53°48’S, 70°53’W to 53°45’S, 70°53'W, 485 m,
Blaketrawl, 6 February 1964, Eltanin Sta. 963 (2 paratypes, USNM 58301).

Description.—All specimens are incomplete; the holotype consists of 58 seti-
gers, is 21 mm long and 1.20 mm wide, with parapodia. The anterior part of the
body (Fig. 3e) is cylindrical and the median and posterior parts are dorsally
flattened. Color patterns and eyes are absent. The prostomium (Fig. 3a) is a short,
rounded lobe (not clearly visible in the illustration) with a pair of triangular frontal

244 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Onuphis (Nothria) lithobiformis (holotype, USNM 58300): a, Anterior end, dorsal view;
b, Pseudocompound hook, third parapodium; c, Pectinate seta, median parapodium; d, Third para-
podium, anterior view; e, Anterior end, lateral view. All scales are 1 mm except where otherwise
indicated.

palps. The ceratophores of the occipital antennae are all strongly ringed and reach
well beyond the edge of the prostomium. The inner lateral ceratophores are the
longest and have up to 13 rings. The outer lateral antennae reach setiger 2, the
inner lateral antennae reach setigers 8—12 and the median antenna reaches setigers
3-7. The outer lateral ceratostyle is clearly shorter than its ceratophore.

Branchiae are present from the first setiger in most specimens. The occurrence
of branchiae on the first 3 setigers may be irregular; in the holotype branchiae
are absent on the second setiger on one side and first present from the third
setiger on the other side. Branchiae are present to the end of all the specimens;
each branchia is simple and straplike and reaches nearly halfway across the dor-
sum in median and posterior setigers.

The first parapodia (Fig. 3d) have rounded acicular lobes; the presetal lobes
follow the outlines of the acicular lobes closely. Distinct contraction folds are
present at the bases of the dorsal cirri. Digitiform postsetal lobes are present in
the first 11-12 setigers. Cirriform ventral cirri are present in the first 4 or 5
setigers; setigers 56 have shortened, blunt cirri; all following setigers have glan-
dular pads replacing the ventral cirri.

VOLUME 95, NUMBER 2 245

Limbate and pectinate setae, pseudocompound and subacicular hooks are pres-
ent. Compound spinigers and large hooks are absent. Limbate setae are present
in all setigers, but are most numerous in anterior setigers. Each of the pectinate
setae (Fig. 3c) is distally oblique and has about 12 teeth; there are 1 or 2 pectinate
setae in a parapodium in median and posterior setigers. Tridentate pseudocom-
pound hooks (Fig. 3b) with short, blunt hoods are present in the first 4 setigers.
The 2 proximal teeth are similar in size and orientation in all hooks; both teeth
are narrowly conical with blunt tips. Bidentate subacicular hooks are present
from setigers 9-15.

The maxillary formula (investigated in one paratype from USNM 67490) is 1+1,
6+8, 440, 6+9 and 1+1; in the unpaired maxilla III the distal tooth is a large
fang, separated from the other 3 teeth by nearly half the length of the maxilla.

The holotype had a flimsy tube, consisting of a thin, fragile inner lining sparsely
covered with sand-grains.

Onuphis (N.) lithobiformis resembles O. (N.) holobranchiata Marenzeller
(1879:132), O. (N.) iridescens (Johnson, 1901:408), O. (N.) opalina (Verrill,
1873:102), and O. (N.) heterodentata, new species (see above) in that all 5 species
have simple branchiae present from the first setiger and have tridentate pseudo-
compound hooks exclusively. The differentiation of O. heterodentata from the
other 4 species was indicated above. Onuphis holobranchiata differs from the
remaining 3 species in that it has the outer lateral ceratophores longer than the
other ceratophores and in that it has digitiform postsetal lobes at least through
setiger 33. The inner lateral ceratophores are the longest in the other 3 species
and none of them have digitiform postsetal lobes in more than 16 setigers. The
middle tooth of the pseudocompound hooks is by far the widest, and is bluntly
conical in O. opalina and O. iridescens; the proximal and the middle tooth are
of the same size and shape in O. lithobiformis, in which no teeth are bluntly
conical.

Etymology.—The general shape of the body resembles that of a centipede,
Lithobius being the name of a well-known genus in that group.

Distribution.—Onuphis lithobiformis is known from upper slope-depths in the
southern Atlantic Ocean.

Onuphis (Onuphis) declivorum, new species
Fig. 4

Material examined.—Atlantic Ocean, off New Jersey, 38°45.2'’N, 73°01.0’W,
350 m, R.V. Pierce Sta. J1-BLM-04B (holotype, USNM 57639).

Description.—The holotype is an incomplete specimen with 39 setigers, is 17
mm long and 1.6 mm wide, with parapodia. The specimen is a female with large
oocytes in the body-cavity. The body is cylindrical throughout and lacks a color
pattern. Eyes are absent.

The prostomium (Fig. 4a) is shorter than wide and has a pair of elongate frontal
palps directed ventrally. The 5 occipital ceratophores are all longer than the length
of the prostomium. The maximal number of rings is 5 and the rings are limited
to the proximal half of each ceratophore. The inner lateral and median cerato-
styles had loosened from the ceratophores, but were retained in a mucus coat
surrounding the specimen. They are all of the same length and would have

246 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

‘ “
a a
| A
a f
3 g
GSN

Fig. 4. Onuphis (Onuphis) declivorum (holotype, USNM 57639): a, Anterior end, dorsal view;
b-d, Inferior, median and superior pseudocompound hooks, second parapodium; e, Second parapo-
dium, anterior view; f, Sixth parapodium, anterior view; g, Ninth parapodium, anterior view; h,
Anterior end, lateral view. All scales are 1 mm except where otherwise indicated.

(o)
jeu

reached about setiger 6, if attached. The outer lateral ceratostyles are about as
long as their ceratophores; the outer lateral antennae reach setiger 2.

The peristomium bears a pair of short, slender peristomial cirri. The first 5
pairs of parapodia (Fig. 4h) are directed forward and slightly ventrally. The para-
podial bases are well developed in the first 3 parapodia and become reduced
posteriorly. In the first several parapodia (Fig. 4e-f) the acicular lobes are evenly
rounded distally and have a contraction fold running across the frontal face at
about the bases of the dorsal cirri. The postsetal lobes are at least as long as the
base of the parapodium and are tapered. Ventral cirri are long and digitiform in

VOLUME 95, NUMBER 2 247

the first 6 parapodia and abruptly replaced by glandular pads from setiger 7. The
dorsal cirri increase in length from the first through the fifth setiger where they
are longer than half the width of the body. From about setiger 10 they remain of
about the same length, but become increasingly slender posteriorly. The postsetal
lobes decrease in size from the first setiger, but are distinct as short, conical
tubercles, even in the last setigers present.

Branchiae are present from setiger 7 to the end of the fragment. The first
branchia is simple; all others are branched (Fig. 4g) with up to 4 long, slender
branchial filaments. The filaments are as long as the dorsal cirri or longer in all
except the first branchial setiger.

Limbate and pectinate setae, pseudocompound and subacicular hooks are pres-
ent. Compound spinigers and large hooks are absent. Tridentate pseudocom-
pound hooks (Fig. 4b-d) with short hoods are present in the first 5 setigers; all
teeth are short and very strongly curved. Bidentate subacicular hooks are present
from setiger 18. Each pectinate seta is distally transverse and has about 15 teeth.
Limbate setae are present in all setigers, but are especially prominent in superior
fascicles in anterior parapodia.

The structure of the maxillae was not observed in the single specimen available.

Onuphis (O.) declivorum resembles O. (O.) acapulcensis Rioja (1944:139), O.
(O.) aucklandensis Augener (1924:418), and O. (O.) vexillaria Moore (1911:266).
All 4 species have branched branchiae starting from an anterior setiger other than
the first and have tridentate pseudocompound hooks exclusively. Of the 4 species,
O. vexillaria has large hooks in some anterior setigers; the other 3 species lack
such hooks. The 3 remaining species can be differentiated on a set of characters.
Onuphis acapulcensis has branchiae from setiger 6 with up to 12 branchial fila-
ments, the occipital ceratophores are strongly ringed, and ventral cirri are cirri-
form in 5 setigers. Onuphis aucklandensis has branchiae from setiger 2 with up
to 5 branchial filaments, the occipital ceratophores are strongly ringed, and ven-
tral cirri are cirriform in the first 7 setigers. Onuphis declivorum has branchiae
from setiger 7 with up to 4 filaments, the occipital ceratophores are poorly ringed,
and ventral cirri are cirriform in the first 6 setigers. These differences are well-
supported by numerical studies on material of 2 of the other species with which
O. declivorum was compared (O. aucklandensis and O. vexillaria).

Etymology.—declivorum, living on the slope, refers to the depth at which the
holotype was collected.

Distribution.—Onuphis declivorum is known from one locality in the Atlantic
Ocean off New Jersey in upper slope-depths.

Onuphis (Onuphis) texana, new species
Fig. 5

Material examined.—Gulf of Mexico, off Brownsville, Texas, 26°16'’29.6’N,
97°10'16.3”W, 13 m, slightly silty sand (87% sand), coll. Thomas Calnan, sta. 243
(holotype, USNM 67491, 2 paratypes, USNM 67492).

Description.—All 3 specimens are incomplete. The holotype consists of 39
setigers, is 10.8 mm long and 0.75 mm wide, with parapodia. The anterior part
of the body is cylindrical, the posterior part is dorsally flattened. The anterolateral
edges of each of the first 10 setigers has a dark brown, nearly black, pigment bar,
visible from both the dorsal and the ventral side. The peristomium has a pair of

248 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

0.05mm

lon

0.5mm

fo)

Fig. 5. Onuphis (Onuphis) texana (holotype, USNM 67491): a, Anterior end, dorsal view; b,
Pseudocompound hook, second parapodium; c, Second parapodium, anterior view; d, Anterior end,
lateral view; e, Pectinate seta, median parapodium. All scales are 1 mm except where otherwise
indicated.

dark brown lateral bars, and each of the ceratophores has 3 equally spaced brown
rounded spots.

The prostomium (Fig. 5a) is narrowed anteriorly and has a pair of ovate frontal
palps. A pair of black eyespots is near the frontal bases of the outer lateral
occipital antennae and another, larger and somewhat diffuse pair of eyespots
(Fig. 5d) is near the outer base of the inner lateral occipital antennae. All cera-
tophores reach well beyond the edge of the prostomium. The inner lateral cera-
tophores are the longest and have up to 18 rings. The styles of the outer lateral
antennae are much shorter than their ceratophores; the outer lateral antennae
reach setiger 2. The inner lateral antennae reach setigers 10-12 and the median
antenna reaches setiger 5 in all 3 specimens.

Branchiae are present from the first setiger to the end of the fragment. The first
18—20 pairs are simple, the next 10 pairs are bifid and, in the last several segments,
either 2 or 3 branchial filaments are found in an irregular arrangement.

The first 2 pairs of parapodia are directed forward. The acicular lobe (Fig. 5c)
is conical and the presetal lobe follows the outline of the acicular lobe closely,
except on the ventral side where it leaves the bases of the setae exposed. The
postsetal lobes are digitiform in the first 10 setigers; thereafter they are reduced

VOLUME 95, NUMBER 2 249

to short, truncate knobs that remain distinct in the remainder of the fragments.
Ventral cirri are cirriform in the first 6 setigers and are replaced by transversely
elongated glandular pads in all setigers thereafter.

Limbate and pectinate setae, pseudocompound hooks, and subacicular hooks
are present. Compound spinigers and large hooks are absent. Limbate setae are
present in all setigers, but are most common in anterior setigers, starting at setiger
3. Median and posterior setigers each have a single, narrow pectinate seta (Fig.
5d) which is distally transverse and has 8-10 teeth. Bidentate pseudocompound
hooks (Fig. 5b) with short, blunt hoods are present in the first 2 setigers only.
Bidentate subacicular hooks are first present from setiger 10.

The maxillary formula (investigated in one paratype) is 1+1, 8+9, 9+0, 5+9
and 1+1.

Tubes were absent.

Onuphis (O.) texana resembles O. (O.) dibranchiata Willey (1905:272) in
having poorly developed branched branchiae present from the first setiger; it
differs in that it has exclusively bidentate pseudocompound hooks; O. (O.) di-
branchiata has both bi- and tridentate kinds. Other species of Onuphis with
branched branchiae present from the first setiger include O. (O.) chinensis Us-
chakov and Wu (1962b:118) and O. (O.) fukianensis Uschakov and Wu
(1962a:93); again both can be separated from O. (O.) texana by having both bi-
and tridentate pseudocompound hooks.

A unique feature of O. (O.) texana appears to be the presence of pseudocom-
pound hooks in only 2 setigers; in all other species of Onuphis such hooks are
present in at least 3 setigers.

Etymology.—tThe species is known from off Brownsville, Texas, hence texana.

Distribution.—The species is known from a single locality in shelf-depths in
the Gulf of Mexico.

Acknowledgments

I would like to thank Mr. Thomas R. Calnan, Bureau of Economic Geology,
University of Texas at Austin, for sending me the interesting material from the
Texan coast and for securing permission to publish the description. The type-
material of Onuphis (Onuphis) declivorum was collected during the offshore in-
vestigations of the Bureau of Land Management, U.S. Department of Interior.
I would like to thank my colleague, Dr. Meredith L. Jones, for advice and for a
critical review of the manuscript.

Literature Cited

Augener, H. 1924. Papers from Dr. Th. Mortensen’s Pacific Expedition 1914-16. X VIII. Polychaeta
II. Polychaeten von Neuseeland I. Errantia.—Videnskabelige Meddelelser fra Dansk Natur-
historisk Forening, Copenhagen 75:241-441.

Fauchald, K. 1980. Onuphidae (Polychaeta) from Belize, Central America, with notes on related
taxa.—Proceedings of the Biological Society of Washington 93(3):797-829.

Hartman, O. 1967. Polychaetous annelids collected by the USNS Eltanin and Staten Island cruises,

chiefly from Antarctic Seas.—Allan Hancock Monographs in Marine Biology 2:1-—387.

, and K. Fauchald. 1971. Deep-water benthic polychaetous annelids off New England to

Bermuda and other North Atlantic areas (Part 2).—Allan Hancock Monographs in Marine

Biology 6:1-327.

250 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Johnson, H. P. 1901. The Polychaeta of the Puget Sound region.—Proceedings of the Boston Society
for Natural History 29:381—437.

Marenzeller, E. von. 1879. Suedjapanische Anneliden I.—Denkschrifte der Mathematisch-Naturwis-
senschaftliche Classe der Kaiserlichen Akademie der Wissenschaften, Wien 41(2):109-152.

Moore, J. P. 1911. The polychaetous annelids dredged by the U.S.S. Albatross off the coast of
southern California in 1904. III. Euphrosynidae to Goniadidae.—Proceedings of the Academy
of Natural Sciences of Philadelphia 63:234—318.

Rioja, E. 1944. Estudios anelidologicos XI. Notas sobre algunas especies de poliquetos de las costas
mexicanas del Pacifico.—Anales Instituto Biologia, México 15:139-145.

Sars, M. 1835. Beskrivelser og lagttagelser over nogle maerkelige eller nye i Havet ved den Bergenske
Kyst levende Dyr af Polypernes, Acalephernes, Radiaternes, Annelidenes og Molluskernes
Classer, med en kort Oversigt over de hidtil af Forfatteren sammesteds fundne Arter og deres
Forekommen. Bergen, xii and 81 pp.

Uschakov, P. V., and B. L. Wu. 1962a. [Preliminary report on the Polychaetes from Chekian and
Fukian, China].—Studia Marine Sinica [Hai Yang K’o Hsueh Chi K’an] 1(1):89-108.

, and . 1962b. [The Polychaetes from the Yellow Sea. VI. Additions to Errantia].—

Studia Marina Sinica [Hai Yang K’o Hsueh Chi K’an] 2(2):110-134.

Verrill, A. E. 1873. Results of recent dredging expeditions on the coast of New England.—American
Journal of Science, New Haven (3) 5:98-106.

Willey, A. 1905. Supplementary report XXX. Report on the Polychaeta collected by Professor Herd-
man, at Ceylon in 1902.—Report to the Government of Ceylon on the Pearl Oyster Fisheries
of the Gulf of Manaar by W. A. Herdman, D.Sc., F.R.S., P.L.S. with Supplementary Reports
upon the Marine Biology of Ceylon by Other Naturalists 4:243-324.

Department of Invertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 251-255

A NEW SPECIES OF THE GENUS CAMBARINCOLA
(CLITELLATA: BRANCHIOBDELLIDA) FROM ILLINOIS
WITH REMARKS ON THE BURSA OF CAMBARINCOLA
VITREUS ELLIS, 1919, AND THE STATUS OF
SATHODRILUS HOLT, 1968

Perry C. Holt

Abstract.—Cambarincola illinoisensis, new species, is described and illustrat-
ed. Similarities between the everted bursa of this species, that of C. vitreus Ellis,
1919, and those of species of Sathodrilus Holt, 1968, are illustrated and com-
mented on. The involved genera are not synonymized, since it is expected future
work will result in the erection of new genera to contain some species now
assigned to each genus.

Recently, in the process of redescribing Cambarincola elevatus Goodnight,
1940, and reassigning the species to Sathodrilus Holt, 1968, specimens of Cam-
barincola vitreus Ellis, 1919, and a previously unrecognized species of Camba-
rincola were found with everted bursae. To describe the newly found species and
comment on the bursae and penes of the 2 species and those of Sathodrilus are
the objectives of this report.

The methods I use have been fully stated (Holt 1960): here it might be empha-
sized that the drawings, made with the aid of a camera lucida, are semi-diagram-
matic, with little or no effort to indicate cellular detail, and those of bursae and
penes are of optical sections done from animals mounted entire. Measurements
are to be regarded as approximations and are given in millimeters with the ranges
in parentheses. The initials “‘USNM7”’ refer to the catalog numbers of specimens
in the collections of the National Museum of Natural History; ‘“‘PCH’’ to those
of specimens in my collection.

I wish to acknowledge the financial aid of the Virginia Polytechnic Institute
and State University; the help given me by Horton H. Hobbs, Jr., and his iden-
tifications of the host crayfishes; the assistance in collecting of my wife, Virgie
F. Holt, and our daughter, Susan E. H. West.

Cambarincola illinoisensis, new species
Fig. 1A—E

Type-specimens.—Holotype, USNM 65225, 3 paratypes, USNM 65226, 5 para-
types, PCH 840, taken on Orconectes virilis (Hagen 1870) from a prairie stream
(? Sugar Creek) north of Stockland, Iroquois County, Illinois, by Perry C. and
Virgie F. Holt, 25 July 1958.

Diagnosis.—Medium-large, relatively robust worms (holotype 3.2 mm in
length); upper lip with 4 obscure lobes, lower with 2; no oral papillae; low dorsal
ridges; upper jaw twice that of lower in size, dental formula 5/4; bursa about 4
body diameter in length, ovate; atrial fold present; protruded penis slender; tu-

252 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Cambarincola illinoisensis: A, B, Holotype; A, Outline of entire animal, lateral view; B,
Lateral view of jaws; C-E, Paratype; C, Lateral view of reproductive systems (b, bursa; p, prostate;
sg, spermiducal gland; spb, spermathecal bulb; spd, spermathecal duct); D, Optical longitudinal sec-
tion through penis and bursa (a, atrium; ab, atrial part of bursa; af, atrial fold; ed, ejaculatory duct;
p, penis; ps, penial sheath); E, Optical longitudinal section through everted bursa and penis. Cam-
barincola gracilis from Lincoln County, Oregon: F, Optical longitudinal section through bursa and
penis. Sathodrilus dorfus from Lincoln County, Oregon: G, Optical longitudinal section through bursa
and penis. Sathodrilus lobatus from Lincoln County, Oregon: H, Optical longitudinal section through
bursa and penis.

VOLUME 95, NUMBER 2 M3

bular spermiducal gland about 14 body diameter in length, and about 1% its length
in diameter; prostate slender, in diameter %4 to 4 that of spermiducal gland,
composed of densely granular cells, with prominent ental bulb; spermatheca
greater than body diameter in length, with long ectal duct, ovate bulb.

Etymology.—For the state of Illinois.

Description.—Five specimens of Cambarincola illinoisensis, including the ho-
lotype, have the following dimensions: total length, 2.9 (2.5-3.3); greatest di-
ameter, 0.5 (0.4—0.6); head length, 0.5 (0.40.7); head diameter, 0.3 (0.3-0.4);
diameter, segment I, 0.3 (0.2-0.4); diameter, sucker, 0.3 (0.2-0.4).

The lobes of the upper lip are very short, broad, and not easily seen in most
specimens; there are no oral papillae. External sulci of the head are shallow and
broad; the one such sulcus other than the peristomial one is a slight ventral
depression. There is one internal pharyngeal sulcus. The dorsal ridges are low
and indistinct, but weakly developed supernumerary segmental muscles are pres-
ent.

The jaws are distinctive: of an irregular triangular shape in lateral view, the
upper bears a proportionately huge median tooth and 2 almost undetectable lateral
teeth on each side; the lower jaw is from ?/s to 12 the upper in length and bears
2 prominent paramedian teeth and 2 minute lateral ones.

The spermiducal gland is not unusual in any respect. It is approximately *4 the
body diameter in length and about % its length in diameter, tapering slightly to
its junction with the ejaculatory duct, and usually lying obliquely in the coelom
of its segment. The vasa deferentia enter the gland at widely separate points,
creating the appearance, from some perspectives, of a prominent anterior deferent
lobe.

The prostate is unusual: the glandular cells which compose it are finely and
densely granular, unlike those of the spermiducal gland which are of the common
type, large and coarsely granular. In shape and size, the prostate is slender, in
diameter about %4 that of the spermiducal gland and subequal to the latter in
length. The ental bulb is prominent, but the prostate itself, lying partly obscured
in lateral view alongside the spermiducal gland, is often difficult to see.

The bursa is elongate ovate, approximately twice its diameter and about 4 the
diameter of the body in length. There is a prominent median atrial fold. The penial
sheath comprises about '4 of the total length of the bursa.

One specimen of the paratype series has an everted bursa (Fig. 1E). The penis
is carried to the outside as a relatively slender tube, but it is impossible to tell if
it itself is everted.

The ejaculatory duct presents no features of note.

The male efferent apparatus as a whole is proportionately small, lying entirely
beneath the gut in some specimens and never extending to the dorsal border of
the gut. The necessity of viewing the structures against a background of the gut
contents makes it difficult to determine the extent and nature of the spermiducal
gland and related structures in whole mounts, the only material available. For-
tunately, and most unusually, in one paratype-specimen the ovarian, VII, segment
iS parasitized by a nematode and, presumably as a result, the spermatozoa that
normally fill the coeloms of segments V and VI are absent and all of the male
secondary sexual organs are easily seen.

The spermatheca is somewhat greater than the body diameter in total length

254 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

and its ectal duct is about twice the length of its bulb which is slender ovate in
shape. There is no ental process.

Variations.—None, other than those associated with size and degree of con-
raction, were noticed. |

Affinities.—Cambarincola illinoisensis seems to be closer to C. jamapaensis
Holt, 1973, than to any other of its congeners. C. illinoisensis is composed of
larger worms. The reproductive systems are of similar proportions, but differ in
the proportionately greater size of the prostate of C. jamapaensis which was
earlier considered as related to species from the Appalachians with differentiated
prostates (Holt 1973:20). C. acudentatus Holt, 1973, also has a prostate composed
of cells that are densely filled with small granules, but differs from C. jamapaensis
and C. illinoisensis in the absence of dorsal ridges and in the shape and size of
its jaws: delicate and light in color with a dental formula of 7/6 (Holt 1973:11-
13). The ‘‘differentiated’’ prostate of these 3 species differ from those of such
Appalachian species as C. philadelphicus (Leidy, 1851) and C. fallax Hoffman,
1963, which have prostates that are composed of heavily vacuolated ceils: those
heretofore described as ‘‘differentiated.’’ The Mexican species, C. jamapaensis,
thus remains for now as the closest relative of C. illinoisensis, but the two are
readily separated by the disparity in the size of the jaws in the latter.

Host.—Orconectes virilis (Hagen, 1870).

Distribution.—Known only from the type-locality; it is expected that C. illi-
noisensis will be found widely throughout the Central Plains of the United States.

Discussion.—As noted, a specimen among the paratypes has an everted bursa.
In addition, 2 specimens of C. vitreus with everted bursae were found in the
material studied for the paper referred to above (Holt 1978). The occurrence of
specimens collected with everted bursae and protruded or everted penes is rare
and in this case furnishes an opportunity to comment further on the structure of
the bursa and penis in the genera Cambarincola and Sathodrilus.

The penis of members of Cambarincola has been considered non-eversible
(Moore 1895:498; Ellis 1912:481; Holt 1949:554; Hoffman 1963:289-—290; inter
alia), while that of species of Sathodrilus has been described as eversible (Holt
1968:294). To my knowledge, no one has actually seen either the bursa or the
penis evert in specimens of these genera and all these statements (certainly mine
and Hoffman’s) are inferences based on the observable structures of dead ani-
mals.

If Cambarincola vitreus and C. illinoisensis were not clearly in other respects
congeneric with the type of the genus (Cambarincola macrodontus Ellis, 1912),
the shape of their almost identical penes carried outward by everted bursae might
align them with species of Sathodrilus. Indeed, a comparison of illustrations of
optical sections, showing in each case the retracted penis (Fig. 1F—H) of repre-
sentative species of these genera does not reveal any significant differences: those
of shape, size, proportionalities of parts and atrial folds are common among the
members of both genera. Species of Sathodrilus differ from those of Cambarin-
cola in lacking prostates, having only what I have called a ‘“‘prostatic protuber-
ance’ (Holt 1968:298), or in having prostates that arise from the spermiducal
gland entad to its junction with the ejaculatory duct.

Taxonomists customarily recognize genera as groups of species separated from
other such similar groups by one or more morphological “‘gaps.’’ Often subse-

VOLUME 95, NUMBER 2 255

quent discoveries or additional studies lead to the obliteration of the ‘‘gaps’’ (Holt
1968:5). Such seems to be the case here: the eversible penis attached by cyto-
plasmic strands to the inner wall of the penial sheath, if indeed this is the correct
description, of some species of Sathodrilus and the size and shape of the prostate
vary through a series of species until the ‘‘gap’’ between Sathodrilus and Cam-
barincola is virtually bridged. However, I do not choose to merge the currently
recognized 44 species assigned to Cambarincola with the 15 that nominally com-
pose the genus Sathodrilus. Rather, I suspect more than 2 genera should be
erected for these 59 species and with a refinement of taxonomic concepts and
procedures in the study of the branchiobdellids, I expect that this will be done.

Literature Cited

Ellis, Max M. 1912. A new discodrilid worm from Colorado.—Proceedings of the United States

National Museum 42(1912):481-486.

. 1919. The branchiobdellid worms in the collections of the United States National Museum,

with descriptions of new genera and new species.—Proceedings of the United States National

Museum 55(2267):241-265, pls. 10-13.

Goodnight, Clarence J. 1940. The Branchiobdellidae (Oligochaeta) of North American crayfishes.—
Illinois Biological Monographs 17(3):1-75, pls. 1-3.

Hoffman, Richard L. 1963. A revision of the North American annelid worms of the genus Camba-
rincola (Oligochaeta: Branchiobdellidae).—Proceedings of the United States National Museum
114(3470):271-371.

Holt, Perry C. 1960. The genus Ceratodrilus Hall (Branchiobdellidae, Oligochaeta) with the descrip-

tion of a new species.—Virginia Journal of Science, new series, 11(2):53—73, pls. 1-4.

. 1968. New genera and species of branchiobdellid worms (Annelida: Clitellata)—Proceedings

of the Biological Society of Washington 81:291-318.

. 1973. A summary of the branchiobdellid worms (Annelida: Clitellata) fauna of Mesoameri-

ca.—Smithsonian Contributions to Zoology 142:i-iii, 140.

. 1978. The reassignment of Cambarincola elevatus Goodnight, 1940 (Clitellata: Branchiob-

dellida) to the genus Sathodrilus Holt, 1968.—Proceedings of the Biological Society of Wash-

ington 91(2):472-482.

Leidy, Joseph. 1851. Contributions to helminthology.—Proceedings of the Academy of Natural Sci-
ence of Philadelphia 5:205—209.

Moore, J. Percy. 1895. The anatomy of Bdellodrilus illuminatus, an American discodrilid.—Journal
of Morphology 10(2):497-540, pls. 28-32.

Department of Biology, Virginia Polytechnic Institute and State University,
Blacksburg, Virginia 24061.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 256-264

A NEW SPECIES OF OREOHELICID LAND SNAIL
FROM THE SAN AGUSTIN PLAINS, NEW MEXICO

Celinda R. Crews and Artie L. Metcalf

Abstract.—A new species of pulmonate land snail, Oreohelix litoralis, is de-
scribed from populations inhabiting wave cut cliffs in the southern part of the San
Agustin Plains, west-central New Mexico. Taxonomic evaluation is based on
qualitative and quantitative characters of shell and genital structures. A rede-
scription of Oreohelix magdalenae Pilsbry is provided, since it is considered to
be closely related to Oreohelix litoralis. Both species are members of the Oreo-
helix metcalfei Cockerell complex within the Oreohelix yavapai Pilsbry group.

Most of the basic taxonomic work on the genus Oreohelix in New Mexico was
done by H. A. Pilsbry and J. H. Ferriss, culminating in Pilsbry’s (1939) mono-
graphic treatment of known species. A number of New Mexican oreohelicids
were assigned to the wide-ranging Oreohelix yavapai Pilsbry, 1905, group. Taxa
occurring in the Black Range and peripheral mountains in south-central New
Mexico were ascribed to the Oreohelix metcalfei Cockerell, 1905, complex. Met-
calf (1974) discussed and referred several species to this complex, including Or-
eohelix magdalenae, Pilsbry, 1939, as a northeastern derivative. The new species
described below is judged to be a northern derivative of this complex and to be
closely related to Oreohelix magdalenae.

Oreohelix litoralis, new species
Figs. 1A, B, C; 2A; 3A

Description of shell of holotype (from Locality 6).—Shell slightly depressed,
convex dorsally and ventrally; spire forming angle of 118°; 16.4 mm in diameter
and 9.7 mm in height; upper lip descending only slightly; angular peripherally
with angularity at about mid-height of body whorl; first 2 whorls keeled; aperture
slightly ovate horizontally, 7.7 mm wide and 6.2 mm high; aperture oriented at
angle of 54° to vertical axis of shell; umbilicus open, not contracting rapidly
within, 3.9 mm wide, contained 4.21 times in diameter; sutures moderately im-
pressed; 5.10 whorls; dorsal surface generally smooth, with sculpture limited to
growth lines, weak and irregularly spaced on early whorls, becoming stronger on
body whorl; ventral surface smooth with only low growth lines (no spiral striae
or lirae); earliest 2.25 whorls of dorsal surface uniformly cinnamon brown, grading
to grayish brown mottling on a whitish background by whorl 3 with mottled
pattern extending to body whorl; thin brown band appears at 2.25 whorls, ex-
tending to body whorl; this band situated approximately one-third whorl-width
inward from suture on earlier whorls and approximately one-third whorl-width
above angulation on body whorl; ventral surface with grayish-brown mottling
mainly descending (“‘bleeding’’) from thin brown band present on body whorl
immediately beneath angulation.

Variation in shells.—Variation in shell measurements and proportions for 20

VOLUME 95, NUMBER 2 Dow

Fig. 1. A, B, C, Shell of holotype of Oreohelix litoralis (diameter, 16.4 mm); D, E, F, Shell of
Oreohelix magdalenae from Loc. 9 (diameter, 15.7 mm).

specimens each from Localities 6 and 7 is given in Table 1. A few paratypes are
less depressed than the holotype. Descent of the body whorl ranges from weak
to marked. Shape of the aperture ranges from round to oval. Banding and other
coloration varies little among paratypes from Localities 1-7. There is a variable
degree of fading, due to wear, on the ventral surface of the last half of the body
whorl.

Description of genitalia (based on dissections of 5 paratypes from Locality 6;
collected in Dec. 1979; see Figs. 2A and 3A).—Lower penis swollen, upper penis
bearing small, rounded appendix laterally; internally, lower penis bears 2-4 ir-
regular longitudinal pilasters, upper penis with close-set pustules, varying in size;

258

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Some shell and genital measurements (mm) and proportions for Oreohelix litoralis and
Oreohelix magdalenae. Mean and standard deviation (in parentheses) are given above and range
below. D = diameter.

Diameter
Height

Aperture
width

Aperture
height

Umbilicus
width

No. of whorls
D/height

D/aperture
width

D/aperture
height

D/umbilicus
width

D/no. of whorls

O. litoralis
Loc. 6; n = 20
18.9 (1.01)
Weil
11.7 (0.89)
10.0-13.6

8.8 (0.64)
7.5-10.3
7.1 (0.42)
6.2-7.8

5.1 (0.47)
4.56.2
5.33 (0.122)
5.10-5.55
1.62 (0.102)
1.38-1.84
2.14 (0.077)
MIP. SII
2.67 (0.135)
2.43-—2.99
3.75 (0.237)
3.384.22
3.55 (0.179)
3.23-4.02

O. litoralis
Loc. 7; n = 20
17.5 (0.86)
16.2-19.9
11.2 (1.03)
10.0-13.6

8.4 (0.75)
7.1-10.4
7.8 (0.57)
6.5-8.8

4.4 (0.39)
3.9=5.1
5.16 (0.141)
4.90-5.40
1.58 (0.096)
| S51 72
2.10 (0.097)
1.91-2.29
2.26 (0.106)
2.05—2.49
4.01 (0.264)
3.49_4.39
3.40 (0.145)
3.11-3.69

O. magdalenae
Loc. 9; n = 20

14.0 (0.88) Penis
285.7

8.5 (0.61) Epiphallus
f-5—95

6.5 (0.38) Vas deferens
S715 1

6.2 (0.29) Vagina
oii)

3.7 (0.30) Free oviduct
3.2-4.3

4.61 (0.206) Spermathecal

4.25-5.00 duct plus sac
1.65 (0.068) D/penis
1.55-1.83

2.15 (0.089) D/epiphallus
2.00—2.31
2.25 (0.136) D/vas deferens
2.02-2.53
3.76 (0.210) D/vagina
3.34-4.11
3.03 (0.112) D/free oviduct
28532
D/spermathe-
cal duct
plus sac

O. litoralis
Loc. 6;n = 5
8.3 (0.41)
7.8-8.9
3.4 (0.41)
2.83.8
6.3 (0.47)
5.8-7.0
3.2 (0.34)
2.9-3.8
3.2 (0.41)
MBL

12.6 (1.03)
We SS
2.00 (0.147)
1562 — es)
4.92 (0.553)
4.42-5.79
2.62 (0.210)
2.39-2.88
5.15 (0.591)
4.42-5.76
5.26 (0.574)
4.67-6.19
1.32 (0.042)
1.26—-1.36

O. magdalenae
Loc. 9; n =
8.9 (0.67)
7.9-9.6
3.4 (0.37)
2.8-3.8
9.8 (0.44)
9.4-10.5
3.2 (0.21)
3.0-3.5
2.7 (0.29)
2.33.0
11.9 (0.72)
10.9-12.6
1.56 (0.106)
1.44-1.69
4.12 (0.379)
3.55—4.61
1.42 (0.083)
1.33—1.51
4.41 (0.366)
4.03-4.77
5.16 (0.606)
4.70-6.22
1.17 (0.077)
1.07—1.27

upper and lower penis approximately equal in length; penial verge short; penial
retractor muscle strands attached to both upper penis and epiphallus at their
juncture; epiphallus relatively long and slender, tapering at upper end; vas def-
erens inserts centrally on upper terminus of epiphallus; vagina stout, widening at
upper end; free oviduct stout, widening at base; albumen gland short and muzzle-
shaped with slightly darker pigmentation; talon hook-like, black in color, slightly
swollen and of moderate size relative to albumen gland; spermathecal sac oval
to suboval. Lengths of some genital organs and ratios of shell diameter to organ
lengths are given in Table 1.

Description of embryonic shells. —Embryos taken from a single dissected spec-
imen from Locality 6 possess 2.55, 2.60, and 2.65 whorls and diameters measure
4.2, 4.1, and 4.2 mm, respectively. Overall color is cinnamon brown as seen in
live adult shells for approximately the earliest 2.5 whorls (first 2.25 whorls in the
holotype). Banding is faint, but discernible dorsally after the second whorl. Em-
bryonic shells are keeled, but possess no cuticular projections.

Remarks.—On 11 October 1980, Oreohelix litoralis was found in crevices and
in rock rubble of wave cut cliffs on the southern margin of the San Agustin Plains
at an elevation of approximately 2105 m (6900 feet) (Loc. 6). Rocks were igneous,

VOLUME 95, NUMBER 2 259

Fig. 2. A, Genitalia of Oreohelix litoralis; B, Genitalia of Oreohelix magdalenae. \p = lower
penis; up = upper penis; ap = appendix; pr = penial retractor muscle; e = epiphallus; vd = vas def-
erens; pg = prostate gland; v = vagina; fo = free oviduct; u = uterus; ag = albumen gland; t = tal-
on; hd = hermaphroditic duct; sd = spermathecal duct; s = spermathecal sac; a = atrium; go =
common gonopore.

mainly rhyolite tuff, welded tuff, and scoria. The area is grassland with sparse
shrubs, in contrast to the forested habitats observed in association with most
occurrences of Oreohelix. Common plants included Atriplex canescens (four-
winged saltbush), Rhus aromatica (lemon sumac), Brickellia sp. (brickellia),
Lycium sp. (wolfberry), and Bouteloua gracilis (blue grama grass).

Live, aestivating individuals were found attached to small rocks in rubble, on
the underside of larger stones in talus, and beneath low ledges at the base of the
cliffs. Oreohelix litoralis at this locality seemed colonial, with up to 11 individuals
clustered under one rock. Both live and dead snails were abundant. The harsher
environment occupied by these populations (relative to montane populations)
may be related to the high degree of colonialism observed.

Disposition of types.—The holotype (Fig. 1A, B, C) is a fresh shell with des-
iccated soft parts (National Museum of Natural History 784661). The paratypes
are fresh shells with desiccated soft parts (National Museum of Natural History
784662; University of Texas at El Paso 7725, 8467, 8593-8594; Academy of Nat-
ural Sciences of Philadelphia 354156; University of Arizona 19016; Dallas Mu-
seum of Natural History 5364).

Etymology.—The epithet litoralis (L., of the shore) refers to occurrence of
populations of the species along wave cut cliffs of the former shoreline of Pluvial
Lake San Agustin.

Oreohelix magdalenae Pilsbry
Figs. 1D, E, F; 2B; 3B

Oreohelix socorroensis magdalenae Pilsbry, 1939: 515, Fig. 336b (Type: Mag-
dalena Mountains, Socorro County, New Mexico; ANSP 158166).
Oreohelix magdalenae.—Metcalf, 1974:99.

260 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. A, Longitudinal dissection of the penis of Oreohelix litoralis; B, Longitudinal dissection
of the penis of Oreohelix magdalenae. pr = penial retractor muscle; e = epiphallus; po = pore; pu =
pustules; pl = longitudinal pilasters; v = vagina; a = atrium; go = common gonopore.

Description of shells (based on 20 specimens from Locality 9).—Shell slightly
depressed to, in a few specimens, pyramidal; diameter 12.8—14.7 mm (mean 14.0
mm); height 7.5-9.5 mm (mean 8.5 mm); descent of body whorl slight to none;
approximately first 2.25 whorls keeled, becoming angular peripherally with an-
gularity at about mid-height of body whorl; aperture rounded to oval perpendic-
ular to parietal wall, 5.9-7.1 mm wide (mean 6.5 mm) and 5.7-6.7 mm high (mean
6.2 mm); umbilicus open and ample, 3.2—4.3 mm wide (mean 3.7 mm), contained
3.34-4.11 times within diameter (mean 3.76 times); 4.25—5.00 whorls (mean 4.61
whorls); dorsal surface generally smooth with faint lirae on earliest approximately
2.5 whorls, not extending to later whorls; transverse growth lines on first 2.5
whorls, becoming more irregular and increasing in strength on outer whorls;
ventral surface sculpture limited to transverse growth lines; dorsally, first 2.5
whorls dark gray, grading to a dull gray and brown mottled pattern intersected
by irregular transverse white bands that increase in area on outer whorls; thin
brown band evident dorsally after first whorl, becoming stronger on later whorls
and extending to body whorl, band located approximately one-third whorl-width
above suture on earlier whorls and one-third whorl-width above peripheral an-
gulation on body whorl; ventrally, irregular gray, white, and brown transverse
zones of coloration with last 0.5 body whorl worn to tan on some specimens;
ventral band located immediately below peripheral angulation, distinct, and ap-
proximately 3 times size of dorsal band. Additional data on variations in shell
measurements and proportions are given in Table 1.

Description of genitalia (based on dissections of 5 specimens from Locality 9,
collected on 20 April 1980; see Figs. 2B and 3B).—Lower penis greatly swollen,

VOLUME 95, NUMBER 2 261

approaching a bulbous condition; large, prominent lateral appendix on upper
penis; internally, lower penis with 2 to 3 irregularly shaped longitudinal pilasters;
upper penis with numerous suboval pustules, varying in size; upper and lower
penis approximately equal in length; penial verge slight; penial retractor muscle
with attachment to both upper penis and epiphallus; epiphallus long and slender;
central insertion of the vas deferens onto the apex of the epiphallus; vagina stout,
slightly swollen at emergence of spermathecal duct; free oviduct also stout, slight-
ly larger at base than top; spermathecal sac oval; albumen gland more darkly
pigmented, relatively small; talon hook-like and black, large relative to albumen
gland.

Of a second group of 5 dissections done on specimens of O. magdalenae col-
lected at Locality 9 on 19 May 1981, 3 specimens of adult size (diameters 14.0,
12.5, and 12.5 mm) had the same overall development of the genitalia as seen in
the first group of dissections. However, 2 seemingly subadult specimens showed
varying degrees of development of the genital organs. One specimen (diameter
10.0 mm) had only slight evidence of the lower genitalia (penis-vagina area only),
with no development of the uterus-prostate area. Another specimen (diameter
10.6 mm) showed the beginning of development of the uterus-prostate area, with
lower genitalia developed, but not of full size.

Lengths of some genital structures and ratios of shell diameter to organ lengths
for 5 adult specimens are given in Table 1.

Remarks.—At Locality 9, O. magdalenae was found living under loose igneous
stones at 2230 m (7320 feet) elevation. Vegetation included Pseudotsuga menziesii
(Douglas fir), Pinus ponderosa (ponderosa pine), and Quercus gambelii (Gam-
bel’s oak) on the slope where collections were made, with Populus angustifolia
(narrowleaf cottonwood) common along a brook below the slope. A few speci-
mens were collected (Robert H. Weber) at another site in the range (Locality 8)
near the summit of North Baldy Peak at approximately 3000 m (9850 feet).

At Locality 9, on 20 April 1980 and 19 May 1981, aestivating individuals oc-
curred under stones both attached and unattached with epiphragms intact. One
individual was active under a stone on 20 April 1980. Two to 3 live individuals
occurred together under some stones.

Embryos were absent in uteri of specimens dissected, indicating that devel-
opment to observable size (or perhaps fertilization) had not yet occurred or that
deposition of young had occurred before these (spring) dates noted. It is logical
to suppose that deposition of young should occur before or during the early phase
of the summer rains to allow subsequent development under favorable conditions.
No detailed account of the life history has been published for any species of
‘Oreohelix in New Mexico.

Distribution

Oreohelix litoralis, as now known, occurs in local populations along the south-
ern margin and on an isolated hill arising from the floor of the San Agustin Plains.
Oreohelix magdalenae occurs in local populations in the Magdalena Mountains,
east of the Plains. These species seem to represent northern, peripheral deriva-
tives of the Oreohelix metcalfei complex. This complex is centered in the Black
Range, to the south, with several peripheral species in outlying ranges (Metcalf

262 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1974:99). Ancestors of the 2 species possibly migrated northward during Pleis-
tocene pluvial episodes. The occurrence at Locality 1 is described by Robert H.
Weber (in litt.) as follows: “an isolated outcrop on the floor of the lake plain,
separated from the cliffs . . . by nearly two miles of open grassland. The hill was
submerged at water levels about 6885 feet [2098 m], and was an island between
that level and a little over 6800 feet [2073 m].’’ Weber (pers. comm.) indicates
that the hill was submerged in Pluvial Lake San Agustin during the late Wisconsin
rise of the lake and that it was, for a time, an island during the time of lake
recession in the earlier Holocene. Thus, colonization of the hill by O. litoralis
seems to have taken place during the past 10,000 years. Mountain ranges im-
mediately south of the San Agustin Plains have not been investigated malacol-
ogically.

Differential Diagnoses and Discussion

Although O. litoralis seems closely related to O. magdalenae, it is judged
sufficiently distinctive to warrant specific rank. Shells of O. litoralis are larger,
overall, than shells from populations of O. magdalenae observed (Table 1), and
possess more strongly indented sutures and a slightly higher spire. In O. litoralis,
no lirae are present either dorsally or ventrally, whereas the dorsal surface of O.
magdalenae possesses faint lirae on the earliest ca. 2.5 whorls. Although a mot-
tled color pattern is observed in both species, shells of O. magdalenae are dom-
inated by a darker gray and brown and exhibit less diffuse mottling, with white
areas being restricted and more distinctly demarcated. Both species possess a
thin brown band dorsally and a second band below the peripheral angulation.
However, in O. litoralis the 2 bands are approximately equal in size with the
lower band diffusing ventrally (“‘bleeding’’), whereas the lower band in O. mag-
dalenae is 3 times the width of the upper and discrete.

The overall shell size and number of whorls observed in O. litoralis and O.
magdalenae generally are closer to those recorded for taxa in the O. metcalfei
complex (Pilsbry 1939:507-514; Metcalf 1974:95—98) than to those observed for
other members of the Oreohelix yavapai group occurring in New Mexico. Both
O. litoralis and O. magdalenae have peripheral angulation at about mid-height
of the body whorl in adult shells. Other geographically peripheral taxa (see above)
of the O. metcalfei complex also are angular peripherally, in contrast to the
carinate condition seen in centrally located (i.e., Black Range) taxa. The periph-
eral taxa of the complex, along with O. litoralis and O. magdalenae, also have
Shells that are less depressed, and with more rounded (biconvex) whorls than
those seen in centrally located members of the O. metcalfei complex. Banding
is relatively prominent and surface sculpture is not elaborate, generally lacking
prominent striae and lirae, in O. litoralis, O. magdalenae, and other pene
taxa of the O. metcalfei complex.

We regard the characters exhibited by O. litoralis, O. magdalenae, and other
peripheral taxa—angulation opposed to strong keeling, higher elevation of the
spire, biconvex shells, relatively smooth surface sculpture, and relative promi-
nence of banding—as conservative and opposed to more advanced characteristics
seen in taxa in the center of distribution (i.e., Black Range) of the Oreohelix
metcalfei complex. These latter species possess varying degrees of carination and

VOLUME 95, NUMBER 2 263

depression of shells, elaborate spiral and/or radial surface sculpturing, and lack
prominent banding. Metcalf (1974:99) noted that banding in O. confragosa Met-
calf, 1974 (which occurs west of the Black Range and which he believed to be
more closely related to the peripheral taxa) is weak in living specimens, but more
prominent in fossil shells. If the two-banded pattern in Oreohelix is an ancestral
feature of the genus in the process of disappearing (Pilsbry 1939:413), then the
evolution towards the loss of banding observed by Metcalf would seem to support
the view that the strongly-banded peripheral taxa of the Oreohelix metcalfei com-
plex are more conservative.

Pilsbry (1939:510) reported the presence of prominent “‘triangular processes’’
(cuticular projections) on the keel and dorsal and ventral surfaces of embryonic
Shells of taxa in the O. metcalfei complex in both peripheral and centrally located
taxa. Unfortunately, embryonic shells of O. magdalenae were not available for
comparison. However, embryonic shells of O. litoralis differ from those reported
for other members of the O. metcalfei complex in possessing a relatively smooth
surface and lacking these cuticular projections.

The genitalia of O. litoralis are similar to those of O. magdalenae in possessing
a swollen lower penis, a lateral appendix situated on the upper penis, and a
relatively reduced albumen gland (Figs. 2A, B). However, the degree of swelling
of the lower penis is much greater in O. magdalenae, approaching a bulbous
condition, and reduction in size of the albumen gland is greater. The appendix
also is larger and more pronounced in specimens of O. magdalenae. Several
subspecies of O. metcalfei were reported by Pilsbry (1939:511) as having the
lower penis “‘much swollen.’’ However, the genitalia of most subspecies have
not been described in detail or illustrated for comparison. No information is
available regarding the presence of a lateral appendix on the upper penis in this
complex, with the exception of O. confragosa (Metcalf 1974:96—98), which pos-
sesses a small, lateral, cornuted appendix. Insertion of the vas deferens is central
in O. litoralis and O. magdalenae, but has not been reported for other taxa of
the O. metcalfei complex.

List of Localities

Localities are listed west to east. Localities 2—7 are in areas of wave-cut cliffs
along the generally southern shore of Pluvial Lake San Agustin.

1 New Mexico: Catron Co.: on isolated outcrop on floor of Pluvial Lake San Agustin (former
island): T6S, Rl4W, SW14 SE%4 NW'4 Sec. 28:2040 m (6700 feet): 33°45’34”N; 108°17'29”W (Robert
H. Weber, collector; Nov. 1981).

2 New Mexico: Catron Co.: T7S, RI4W, NW4%4 NE%4 NW Sec. 2: 1950 m (6400 feet): 33°44’05’N;
108°15'26”W (Robert H. Weber, collector; Sept. 1981).

3 New Mexico: Catron Co.: T6S, RI4W, NE%4 NE% SW Sec. 26: 1950 m (6400 feet): 33°45’26’N;
108°15'22”W (Robert H. Weber, collector; Sept. 1981).

4 New Mexico: Catron Co.: shore of Pluvial Lake San Agustin: T6S, R14W, NE%4 SE%4 SW Sec.
35: 2040 m (6700 feet): 33°44'24"N; 108°15'18”W (Robert H. Weber, collector; Sept. 1981).

5 New Mexico: Catron Co.: T6S, RI4W, SE%4 SE™%4 SE'% Sec. 24: 2075 m (6800 feet): 33°46’00’N;
108°13’44”W (Robert H. Weber, collector; Sept. 1981).

6 Type locality of Oreohelix litoralis. New Mexico: Catron Co.: center, boundary T6S, R11W, Sec.
6 and T6S, R12W, Sec. 1: 2105 m (6900 feet): 33°48’54"N; 108°01'16”W (collected by Robert H. Weber,
Dec. 1979, and by A. L. Metcalf and C. R. Crews, 11 Oct. 1980).

264 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

7 New Mexico: Catron Co.: T5S, RILW, SW44 NWi4%4 SW'4 Sec. 27: 2105 m (6900 feet): 33°50'28’N;
107°59'01"W (Robert H. Weber, collector; 28 Nov. 1979).

8 New Mexico: Socorro Co.: Cibola Natl. Forest: Magdalena Mts.: summit of North Baldy Peak:
T3S, R3W, SE%4 SW'4 NW% Sec. 17: 3005 m (9858 feet): 34°03'02”N; 107°10'56”W (Robert H. Weber,
collector; May 1976).

9 New Mexico: Socorro Co.: Cibola Natl. Forest: Magdalena Mts.: T3S, R3W, SW'4 SE'%4 Sec.
21: North Fork Cn.: 2.6 km (1.6 mile) by road from Water Cn.: base of N-facing slope: 2230 m (7320
feet): 34°01'47’N; 107°09'16"W (A. L. Metcalf and C. R. Crews, collectors; 20 April 1980; 19 May
1981).

Acknowledgments

We are grateful to Dr. Robert H. Weber, New Mexico Bureau of Mines and
Mineral Resources, who first apprised us of Oreohelix litoralis and generously
supplied specimens from several localities.

Literature Cited

Metcalf, A. L. 1974. Peripheral species of the Oreohelix metcalfei Cockerell complex (Pulmonata:
Oreohelicidae).—The Nautilus 88:94—100.

Pilsbry, H. A. 1939. Land Mollusca of North America (north of Mexico).—Academy of Natural
Sciences, Philadelphia, Monograph 3:1(1):i-xvii + 1-573 + i-ix.

Laboratory for Environmental Biology, University of Texas at El Paso, El
Paso, Texas 79968.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 265-268

A NEW TREE SNAIL, GENUS DRYMAEUS (BULIMULIDAE)
FROM SOUTHEASTERN PERU

Fred G. Thompson and Jane E. Deisler

Abstract.—Drymaeus aurantiostomus, n. sp. is described from Madre de Dios
Province, Peru. It belongs to the D. expansus species group and is the southern-
most member of the group.

In this paper we describe a new species of tree snail belonging to a group that
is widely distributed in the Andean region of northwestern South America and
adjacent Panama. The snail comes from the Department of Madre de Dios in
southeastern Peru and is the southernmost member of the species group to which
it belongs. The molluscan fauna of this department is very poorly known. No
paper deals specifically with the region. We are grateful to Roy W. McDiarmid,
Department of Amphibians and Reptiles, National Museum of Natural History
for sending us the snail for study and to Alan Solem, Field Museum of Natural
History (FMNH) for the loan of comparative specimens.

Drymaeus aurantiostomus new species
Rigs. i, 2,.5,.6

Diagnosis.—A species of the subgenus Drymaeus Albers, and of the expansus
Species group because of its broadly expanded peristome, it has a relatively ru-
gose axial sculpture and a compressed imperforate umbilicus. It differs from
related species by its color. It is white with a subperipheral and basal spiral band,
and a spiral series of rust colored flames above each band. The interior of the
peristome is bright yellow, thus the Latin derivation of the specific name auran-
tiostomus.

Shell (Figs. 1, 2).—Medium sized, about 30 mm long; conical with a broadly
reflected peristome. Thin, weakly translucent. Spire nearly straight-sided. Um-
bilicus compressed-rimate. Whorls 5.8, nearly flat-sided with a distinct but weakly
impressed subperipheral suture. Whorls regularly increasing in size and descent;
insertion of peristome rising onto periphery of previous whorl; 2.1 embryonic
whorls. Post embryonic whorls sculptured with distinct, regularly spaced, axial
riblets that are crossed by much finer incised spiral striations; axial sculpture
strongest on last whorl behind lip. Peristome broadly expanded throughout, con-
spicuously dilated baso-laterally. Aperture oblique in lateral profile, lying at about
22° to shell axis, nearly tangential to body whorl. Columellar lip broadly expanded
in front of umbilical area. Columella straight, nearly vertical. Color: Embryonic
whorls grayish yellow; postembryonic whorls white with a subperipheral and a
basal purple band on last whorl; each band bordered above by regularly spaced
rusty flames; those above subperipheral band continuing on to penultimate whorl
as a series of small reddish brown spots (Fig. 2); interior of aperture white with
outer bands and flames clearly distinguishable. Peristome with a broad yellow
zone that continues onto columella; edge of peristome white. Parietal callus thin,
clear, colorless.

266 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-4. Drymaeus spp.: 1, 2, D. aurantiostomus, holotype; 3, 4, D. latitesta, holotype.

Measurements of holotype: shell height, 29.2 mm; shell width, 19.7 mm; ap-
erture height 17.1 mm; aperture width, 16.0 mm.

Radula.—Lost during specimen preparation.

Reproductive system (Figs. 5, 6).—Ovotestis large, occupying nearly entire
digestive gland and consisting of numerous closely packed alveolae. Hermaph-
roditic duct highly convoluted; lower half greatly swollen. Albumen gland 8.9 mm
long, lying completely beneath right side of stomach, crescent-shaped with a deep
transverse intestinal groove across distal end. Talon 3.6 mm long, club-shaped,
flexed in middle, completely exposed. Carrefour lying along columellar side of
albumen gland. Uterus about 30 mm long, voluminous and highly convoluted.
Spermatheca globose, lying along columellar side of uterus beneath pericardium
and just below junction of uterus and albumen gland. Spermathecal duct 23 mm
long, tapered. Vagina 5.3 mm long, moderately voluminous. Atrium short. Penis
about 7 mm long (severed during dissection but apparently recovered entire).
Penis simple, slender, with 5 longitudinal branching pilasters internally; demar-
cated from epiphallus by slight internal constriction of lumen. Base of penis with
a short thick penis sheath about 2 mm long. Epiphallus 7 mm long, slender,
cylindrical, terminated by an epiphallic flagellum 1.6 mm long. Lumen of epi-
phallus with low, narrow tubercles anastomosing in lower portion; upper portion
with 5—6 longitudinal branching pilasters that continue diminished into flagellum.
Penis retractor muscle short, 3 mm, inserting on inner wall of lung about % of
distance from pericardium to mantle collar; originating on apex of flagellum. Vas
deferens emerging at junction of epiphallus and flagellum, completely embedded
in epiphallic and vaginal wall; partially exposed above penis sheath.

Type-locality.—Peru, Dept. Madre de Dios, Tambopata Reserve, on Rio Tam-
bopata at Rio La Torre, about 30 km SSW Puerto Maldonado (12°49’ S, 69°17’ W);
280 m alt. Holotype: UF 26605; collected 19 November 1979 by Roy W. Mc-
Diarmid on a shrub along the Tres Chimbadas Trail, about 20 minutes from the
Explorers Inn.

Discussion.—The Bulimulidae are anatomically conservative. Subfamilies are
weakly differentiated. Genera and subgenera usually differ only by shell char-
acters, and occasionally by slight anatomical features. Studies on Drymaeus
anatomy have failed to find anatomical traits that are useful for group classifi-
cation (Strebel 1882; Baker 1925; Pilsbry 1946; Solem 1955; van Mol 1971; Breure
1976; Breure and Eskens 1977). Breure (1979) demonstrated that the digestive
and reproductive systems in Drymaeus are highly conservative. Minor variations

VOLUME 95, NUMBER 2 267

intestinal groove

ovotestis

albumen gland

__ albumen gland

penis retractor ene a!
talon

spermatheca i + e hermaphraditic duct
fi vs ; ay
_/ _epiphallus ge Cre
S — y—— prostate

uterus

\
S— flagellum

-—— prostate

Za

vas deferens

_!-penis sheath

atrium

Figs. 5-6. Drymaeus aurantiostomus, holotype: 5, Reproductive system; 6, Upper part of female
system showing exposed talon. Scales in mm.

in the spermatheca, penis complex and radula have evolved independently in
different species groups and have little phylogenetic significance, although they
may be useful for comparing closely related species. It is apparent that in Dry-
maeus the shell is an adequate indicator of group relationships and specific dis-
tinctions. Data from other morphological systems have not provided a better
basis for classification. The reproductive system of D. aurantiostomus is de-
scribed so that other species may be compared with it. However, other members
of the expansus group have not been studied anatomically.

D. aurantiostomus is related to a group of Andean species with a broadly
expanded peristome and relatively rugose axial sculpture. The group includes D.
expansus (Pfeiffer, 1848), and its subspecies vanattai Pilsbry, 1898, balboa Pils-
bry, 1926, subprotractus Pilsbry, 1902, flavilabrum Weyrauch, 1967, orcesi Wey-
rauch, 1958, altorum Weyrauch, 1958 and pereninus DaCosta, 1901, as well as
D. inca Smith, 1943, D. latitesta Haas, 1952, D. protractus (Pfeiffer, 1885), D.
bartletti (Adams, 1866), D. scitus (Adams, 1885), D. eusterius Pilsbry, 1944, and
D. weeksi Pilsbry, 1926. The group is widely distributed in the northern Andean
region in Panama, Columbia, Ecuador, and Peru. D. aurantiostomus and D.
latitesta are the southernmost members of the group and come from adjacent
areas in Peru.

D. aurantiostomus is similar in shape and size to D. expansus. D. expansus
differs by having more rugose sculpture, the peristome is not as protracted baso-
laterally, the color pattern consist of purplish flammulate blotches on a grayish
white background, a basal band is absent, the interior of the aperture is purple
tinted, and a subperipheral band is usually absent, except in some specimens in
which the flames may fuse to form a band.

268 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

D. aurantiostomus is similar to D. latitesta Haas, a species described from
Dept. Cuzco, Peru. The holotype of D. latitesta (FMNH 38120) (Figs. 3, 4) is a
partly weathered specimen with a chipped peristome and broken apex, but im-
portant characteristics of shape and color pattern are clear. They are alike in
having a subperipheral band that enters the aperture below the insertion of the
outer lip (the band is faded on the front of the holotype of D. latitesta) and in
having a basal band that enters the aperture at the insertion of the columellar lip.
Both species have flammulate markings above the bands, though they are better
defined and more periodic in D. aurantiostomus. D. latitesta differs from D.
aurantiostomus in structure by having more inflated whorls with a more deeply
impressed suture. It is larger (37.2 mm long), is more corpulent in shape, and the
axial striations are weaker (Fig. 4). The columella is nearly vertical and the outer
peristome is nearly uniformly arched. The plane of the aperture in lateral profile
is vertical and basal in position, not tangential to the body whorl. The color
pattern differs from that of D. aurantiostomus by having an additional supraper-
ipheral band with flames above it (Fig. 4).

Literature Cited

Baker, H. B. 1925. The Mollusca collected by the University of Michigan-Williamson Expedition in
Venezuela. IV.—Occasional Papers of the Museum of Zoology University of Michigan 167:1-
49, pls. 12-19.

Breure, A. S. H. 1976. Notes on Bulimulidae (Gastropoda, Euthyneura): Some Bulimulidae from

French Guyana and Surinam with notes on their anatomy.—Zoologische Mededelingen Rijks-

museum Natuurlijke Historie Leiden 50:107-115.

. 1979. Systematics, phylogeny and zoogeography of Bulimulidae (Mollusca).—Zoologische

Mededelingen Rijksmuseum Natuurlijke Historie Leiden (168): 1-215, pls. 1-3.

, and A. A. C. Eskens. 1977. Observations on the formation of spermatophores in a bulimulid

land snail, Drymaeus canaliculatus (Pfeiffer, 1845) (Mollusca, Gastropoda, Pulmonata).—

Netherlands Journal of Zoology 27: 271-276.

van Mol, J.-J. 1971. Notes anatomiques sur les Bulimulidae (Mollusques, Gastropodes, Pulmones).—
Annales de la Société Royale Zoologique de Belgique 101:183—225.

Pilsbry, H. A. 1946. Land Mollusca of North America (north of Mexico).—Monographs (3) Academy
of Natural Sciences of Philadelphia 2(1): 1-520.

Solem, A. 1955. Mexican molluscs collected for Dr. Bryant Walker in 1926, IX. Drymaeus.—Oc-
casional Papers of the Museum of Zoology University of Michigan (566): 1-20, pls. 1-5.
Strebel, H. 1882. Beitrag zur Kenntniss der Fauna mexikanischer Land—und Susswasser-Conchy-

lien. Pt. 5. Orthalicidae. 1-144, pls. 1-19—Hamburg.

Florida State Museum, University of Florida, Gainesville, Florida 32611.

PROC. BIOL. SOC. WASH.
95(2), 1982, p. 269

YOCHELSONIELLA, NOM. NOV., A NEW NAME FOR
ELLISELLA ROHR, 1980 (GASTROPODA)
NON GRAY, 1858 (COELENTERATA)

David M. Rohr and Richard W. Huddleston

Rohr (1980) erected the new genus Ellisella (type-species E. greggi Rohr, 1980)
for Middle-Upper Ordovician Archaeogastropoda from the Kangaroo Creek For-
mation, California. Rohr (1980) tentatively assigned this form to the family Euom-
phalidae. Gray (1858) previously used the name Ellisella for Recent anthozoan
coelenterates. This renders Ellisella Rohr, 1980 a junior homonym of Ellisella
Gray, 1858 and requires a new name (International Code of Zoological Nomen-
clature Article 53, 60a).

We propose the new name Yochelsoniella to replace Ellisella Rohr, 1980 non
Gray, 1858.

Yochelsoniella is characterized by possessing ‘“‘Nearly flat-spired, rounded
nodes at upper-outer angulation; small rounded tubes at mid-whorl; subovate
whorl section; widely umbilicate’’ (Rohr 1980:166).

Yochelsoniella is constructed in honor of Dr. Ellis Yochelson for his contri-
butions in Paleontology.

Acknowledgments

We thank Chevron Oil Field Research Company for their assistance.

Literature Cited

Gray, J. E. 1858. Synopsis of the families and genera of axiferous zoophytes or barked corals.—
Proceedings of the Zoological Society of London 25 (1857):278-294.

Rohr, D. M. 1980. Ordovician-Devonian Gastropoda from the Klamath Mountains, California.—
Palaeontographica Abteilung A 171:141-199.

(DMR) Department of Geology, Sul Ross State University, Alpine, Texas
79830; (RWH) Chevron Oil Field Research Company, La Habra, California 90632
(current address: Scientific Research Systems, 11044 McGirk, El Monte, Cali-
fornia 91731).

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 270-281

PYCNOGONIDA OF THE WESTERN PACIFIC ISLANDS
I. THE MARSHALL ISLANDS

C. Allan Child

Abstract.—This first report on Pycnogonida from the Marshall Islands lists 6
shallow water species. These are: Ammothella stauromata, new species; Ano-
plodactylus glandulifer Stock, previously known from the Red Sea to Singapore;
Anoplodactylus marshallensis, new species; Endeis nodosa Hilton, known only
from Hawaii in past reports, with a male figured for the first time; specimens very
near Callipallene novaezealandiae (Thomson), a species not known from the
western north Pacific; and Nymphon micronesicum, new species. The new
species are apparently not closely related to Pacific mainland pycnogonids.

There are no previous reports on pycnogonids from the Marshall Islands and,
indeed, practically nothing is known of the pycnogonids inhabiting the vast ex-
panse of Micronesia. Only 2 Micronesian species are reported in the literature,
each from a single locality in the Caroline Islands (Stock 1968:10, 49). Tropical
Pacific island pycnogonids have received some study in limited localities by Lo-
man (1908) for Indonesia, Stock (1953) for Indonesia and the Philippines, Hilton
(1942a) for Hawaii, and Stock (1968) for several Pacific localities, with a few short
reports by others (Child 1970, Society Islands and Tuamotus; Child and Hedgpeth
1971, Galapagos; Clark 1973, New Britain and Tonga; Child 1977, French Ocean-
ia). The pycnogonid fauna of Australia, New Zealand, and continental Pacific
borders is better known, probably because it is more accessible to collectors.

The Micronesian fauna could be expected to contain species common to other
parts of the tropical Pacific and at least 3 of the Marshall Island species reported
here are found elsewhere in the Pacific. The remaining 3 species are new, reflect-
ing the lack of benthic collections from Micronesia. The atolls and small volcanic
islands of Micronesia are of particular interest to the taxonomist because of the
restriction or absence of habitats such as muddy estuaries, algal encrusted rocky
shores, and beds of sea grasses. These restrictions limit many marine faunules,
including pycnogonids, to coral or coral sand and rubble, sponges, and a few
other sessile sources of food. Coelenterates are a preferred food of many pyc-
nogonids, many of which have been collected in association with coral reefs. I
predict that there will be a much larger pycnogonid fauna found associated with
the reefs of the Marshall Islands and other Pacific islands and atolls than is
described in this short report.

The majority of these specimens were collected by the author during a Smith-
sonian Enewetak Atoll survey in 1969. Additional specimens were loaned by the
B. P. Bishop Museum of Honolulu, Hawaii.

I am grateful for field support and transportation supplied by the Atomic Energy
Commission who operated the Enewetak Marine Biological Laboratory under the
management of the University of Hawaii, for base support supplied by Kentron,
Inc., and the U.S. Air Force, and for the field support of my fellow divers from
the Smithsonian and the U.S. Geological Survey. I wish to thank Dr. Dennis M.

VOLUME 95, NUMBER 2 271

Devaney of the B. P. Bishop Museum for loaning the additional specimens. The
Specimens marked (BPBM) are deposited in the Bishop Museum. All other spec-
imens are deposited in the National Museum of Natural History under the catalog
numbers of the old U.S. National Museum (USNM). I wish to thank Dr. Thomas
E. Bowman, Department of Invertebrate Zoology (Crustacea), National Museum
of Natural History, for critically reviewing the manuscript.

Family Ammotheidae
Ammothella stauromata, new species
Fig. 1

Material examined.—N end of Enewetak Island, on pilings of large marine pier
in 0.3-1.8 m, 11°21'48’N, 162°21'10’E, Child sta. 39-69, 12 Oct. 1969, 1 male
holotype, USNM 189275, and 6 juvenile paratypes, USNM 189276. Pilings of
small wooden pier at N end of Enewetak Island in 1.5 m, 11°21'52’"N, 162°21'15’E,
Child sta. 25-69, 29 Sept. 1969, 7 juvenile paratypes, USNM 189277.

Description.—Moderately small, adult leg span just under 14 mm. Trunk As-
corhynchus-like with slender median tubercles taller than their segment diameter.
Trunk segmentation flaring with each segment anterior inserted into posterior
large cowl of each preceding segment. Lateral processes separated by distances
slightly greater than their diameters, twice as long as their maximum diameters,
and armed with short slender dorsodistal tubercles and lateral setae shorter than
segment diameter. Lateral setae arranged only on posterior 6 lateral processes:
1 on posterior of second lateral processes; 1 on anterior and posterior of third
and fourth processes. First lateral processes without setae. Ocular tubercle a
cylinder as tall as median trunk tubercles, placed at anterior of cephalic segment,
with darkly pigmented eyes at rounded tip. Abdomen cylindrical, slightly swollen
distally, longer than median tubercles, carried almost erect, armed with several
short distal setae.

Proboscis moderately inflated with broad flat lips just distal to slight constric-
tion.

Chelifores 3-segmented, slender, with small ovoid vestigial chelae having only
hint of fingers. First segment short, with slender dorso-distal tubercle longer than
segment diameter, without setae. Second segment 2.5 times length of first, with-
out tubercles but with fringe of distal setae as long or longer than segment di-
ameter.

Palps 9-segmented, originating from bulbous tubercles placed anterolaterally
on cephalic segment ventral to ocular tubercle. Each bulbous tubercle with a
slender, short, obliquely-pointing tubercle. Palp slender, second and fourth seg-
ments with single long lateral seta and several short setae. Distal segments with
many short ventral and lateral setae.

Oviger 10-segmented, originating ventral to first lateral processes. First 5 seg-
ments with few short setae, sixth with 8 or 9 short setae, seventh with 2 long
lateral setae, and terminal 3 segments (strigilis) with 2 denticulate spines on each
segment. Spines with many serrations per side.

Third leg: first coxa with slender dorsodistal tubercle not as long as segment
diameter, with shorter tubercle anteriolateral to this, segment armed with ante-
rior, posterior and ventral setae. Coxa 2 with | dorsal seta and several short distal
setae, with slender ventrodistal sexual pore tubercle having several distal setae.

DAD PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Ammothella stauromata, holotype, male: a, Trunk, dorsal view; b, Trunk, lateral view;
c, Third leg with enlargement of cement gland; d, Terminal segments of third leg, enlarged; e, Palp;
f, Oviger; g, Terminal segments of oviger, enlarged; h, Juvenile chelafore.

VOLUME 95, NUMBER 2 2H,

Tubercle almost as long as segment diameter. Coxa 3 with several ventral setae.
Femur armed with single long anterior and posterior midlateral setae, several
ventral short setae, fringe of distal setae, and dorsodistal broad tubercle carrying
cement gland tube along its dorsal length. Tibia 1 the longest segment, tibia 2
shorter than femur, both armed with few dorsal and lateral setae longer than
segment diameter and several shorter ventral setae. Tarsus very short, triangular,
with | dorsal seta and 4 or 5 ventral short setae. Propodus robust, strongly curved,
armed with 3 stout heel spines, 10 or 11 short sole spines, several dorsal setae,
the longest slightly longer than segment diameter, and a fringe of distal setae.
Claw moderately curved, about 0.4 length of propodus. Auxiliary claws almost
as long as main claw.

Measurements of holotype (in mm).—Trunk length (tip of ocular tubercle to tip
4th lateral processes), 1.74; trunk width (across 2nd lateral processes), 1.06; pro-
boscis (lateral), 1.23; abdomen (lateral), 0.71; 3rd leg, coxa 1, 0.32; coxa 2, 0.69;
coxa 3, 0.47; femur, 1.35; tibia 1, 1.42; tibia 2, 1.25; tarsus, 0.18; propodus, 0.68;
claw, 0.27.

Distribution.—Known only from the type-locality, Enewetak Island, Enewetak
Atoll, in the intertidal.

Etymology.—The species name is Greek and means a palisade or stockade,
this pertaining to the palisaded appearance of the tall ocular tubercle, median
trunk tubercles and abdomen as seen in lateral view.

Remarks.—There are few Ammothella species with tall median trunk tubercles.
Those that appear to be closest to this new species are Ammothella setosa Hilton,
1942b, A. menziesi Hedgpeth, 1951, A. thetidis Clark, 1963, and A. exornata
Stock, 1975b. Of these, A. stauromata is least like A. exornata, a very small
compact species with many coxae, chelifore and lateral process tubercles and a
tuberculate ocular base. The median trunk tubercles of A. menziesi are shorter
than the ocular tubercle and are blunt finger-like projections, unlike those of A.
stauromata. Hedgpeth’s species is also notably larger than A. stauromata. The
2 species most similar to this new species are A. setosa and A. thetidis. They are
both slender graceful species with lateral processes well separated, but unlike A.
stauromata, neither has a median trunk tubercle on the cephalic segment and
their chelifores are much longer and more slender.

Family Phoxichilidiidae
Anoplodactylus glandulifer Stock

Anoplodactylus spec., Calman, 1923:289.
Anoplodactylus glandulifer Stock, 1954:80-84, fig. 36; 1958:3; 1968:49; 1974: 16—
17.—Arnaud, 1973:955, figs. 1, 2.

Material examined.—N end of Enewetak Island, pier pilings of large marine
pier in 0.3-1.8 m, 11°21'48’”N, 162°21'10"E, Child sta. 39-69, 12 Oct. 1969, 1 3
with eggs.

Remarks.—This single male (a larval specimen accompanied the male, but it
is too juvenile for determination and may not be this species) is slightly smaller
than the measurements given by Stock (1954:84). It varies in other small details:
the lateral processes are more closely spaced; there are fewer teeth on the chela

274 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

fingers; there are only very low palp buds with no discernable segmentation line;
the propodal lamina is shorter, but is still longer than half the sole length; and
the second oviger segment is longer than that shown for the type. There are also
3 cement glands on each femur of all legs. Stock found either 2 or 3 glands on
legs of the type. This egg-carrying specimen is otherwise the same as the type-
specimen, also a male.

The known distribution of this species, previously from the Red Sea and east
coast of Africa to Burma and Singapore in depths from littoral to 5 meters, is now
extended to the Marshall Islands littoral.

Anoplodactylus marshallensis, new species
Fig. 2a-f

Material examined.—N end of Enewetak Island, on pier pilings of large marine
pier in 0.3-1.8 m, 11°21'48’N., 162°21'10"E., Child sta. 39-69, 12 Oct. 1969, 1
6 with eggs, holotype (USNM 189278), 2 d, 1 2 ovigerous, 1 2, 4 juveniles,
paratypes (USNM 189279). N end of Enewetak Island, pilings of small wooden
pier in 1.5 m, 11°21’52”N, 162°21'15”E, Child sta. 25-69, 29 Sept. 1969, 1 ¢ with
eggs, | larva, paratypes (USNM 189280).

Description.—Very small, leg span only slightly over 4 mm. Trunk robust,
ovoid in dorsal aspect, unsegmented. Lateral processes glabrous, separated by
distances equal to half their diameter, each only slightly longer than its diameter.
Ocular tubercle a moderately tall cone inflated at midlength with unpigmented
eyes, with lateral papillae distal to eyes. Neck fairly short, without setae. Ab-
domen slightly shorter than ocular tubercle, curved dorsally, armed with 3 or 4
distal setae.

Proboscis a cylinder tapering distally to rounded lips, without constrictions.

Palps entirely lacking.

Chelifores robust, club-shaped, curved ventrally in lateral aspect, armed with
1 or 2 dorsodistal setae. Chela tiny, palm ovoid, longer than fingers, armed with
3 or 4 short distal setae. Immovable finger slightly curved, with 1 seta, movable
finger with greater curve, crossing immovable finger at tip, armed with 3 setae.
Fingers without teeth.

Oviger moderately short, second segment curved, third straight and the anes
segment, both armed with several short setae. Fourth segment only slightly longer
than fifth, sixth round, no longer than wide. Fifth and sixth armed with several
setae longer than segment diameters.

Legs moderately short, armed with very few setae and a very long dorsodistal
seta on each major segment. Femur the longest segment, with tibia 1 longer than
tibia 2. Single cement gland a flask-shaped distally-pointing tube shorter than
segment diameter. Body of tube creased with many circumferential constrictions
when seen under high magnification. Tarsus very short, almost triangular, armed
with 3 or 4 short ventral setae. Propodus robust, almost straight, with marked
heel bearing 1 stout spine and 4 short setae. Sole armed with 5 or 6 short curved
spines, a very short lamina and several lateral and distal setae. Claw moderately
curved, robust, two-thirds propodal length. Auxiliary claws entirely lacking.

Measurements of holotype (in mm).—Trunk length (chelifore insertion to tip
4th lateral processes), 0.68; trunk width (across Ist lateral processes), 0.45; pro-

VOLUME 95, NUMBER 2 Zi)

boscis length (lateral), 0.28; abdomen length (lateral), 0.14; 3rd leg, coxa 1, 0.14;
coxa 2, 0.24: coxa 3, 0.14; femur, 0.38; tibia 1, 0.33; tibia 2, 0.28: tarsus, 0.06;
propodus, 0.25; claw, 0.17.

Distribution.—Known only from the type-locality, Enewetak Island, Enewetak
Atoll, in littoral depths.

Etymology.—Named for the Marshall Islands, its type-locality.

Color notes. —When freshly collected, all 11 specimens displayed a distinctive
red color pattern. The white trunk bore a purplish red longitudinal stripe mid-
dorsally from the abdomen tip to the posterior surface of the ocular tubercle.
This stripe was bordered in darker red and was a dark blood-red at the ocular
tubercle. The tibiae each had the same dorsal coloration with tibia 2 slightly
darker. The remainder of the legs and all other appendages were white. After
several months in alcohol, these colors had entirely disappeared. The coloration
appeared to be in the chitinous integument itself and not associated with the gut
diverticula where coloration is more often found among the pycnogonids.

Remarks.—This small species is without many distinctive characters except
for the coloration described above. It appears to be morphologically related to
Anoplodactylus quadratispinosus Hedgpeth, 1943, and to A. inermis Losina-Lo-
sinsky, 1961. Both share with the new species the characters of a tall conical
ocular tubercle, a distinctive large heel on the propodus, and a trunk with lateral
processes separated by distances equal to or less than their diameter. Several
other characters immediately separate this new species from the others. It has
much shorter leg segments, shorter and more robust chelifores, a smaller pro-
boscis, shorter lateral processes, and an unsegmented trunk. Losina-Losinsky’s
species is only known from females, but the characters above would serve equally
to separate A. marshallensis from a male of A. inermis.

The small robust shape of this new species is so far unique to the genus An-
oplodactylus from the western Pacific.

Family Endeidae
Endeis nodosa Hilton
Fig. 2g-1

Endeis (Phoxichilus) nodosa Hilton, 1942a:47-48, fig. 4.
Endeis nodosa.—Stock, 1968:59 [key].

Material examined.—Runit (site name Yvonne) Island, pier pilings on lagoon
side in 0-0.7 m, 11°32'40’N, 162°21'53”E, Child sta. 36-69, 8 Oct. 1969, 2 °
(USNM). Enewetak Island, on lagoon rocks in 4 meters, J. Lamberson, Jan.
1976, 1 2 (BPBM). Hawaii, Honolulu Harbor on boat hawser, O. Degener, Apr.
1947, 1 6 with eggs (USNM).

Remarks.—This species has never been adequately figured and since the male
has never been recorded in the literature, I include a set of figures here of the
Honolulu male specimen. The species is rather distinctive in a genus with many
species difficult to separate. The intestinal diverticula in each leg have many blind
caeca or pockets and the femur has a large ventral triangular projection on each
leg. Endeis flaccida Calman, 1923, is a similar species, but is without the ventral
_ femoral projection.

276 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Anoplodactylus marshallensis, holotype, male; a, Trunk, dorsal view; b, Trunk, lateral
view; c, Chela; d, Third leg with enlargement of cement gland; e, Terminal segments of third leg,
enlarged; f, Oviger. Endeis nodosa, male: g, Trunk, dorsal view; h, Third leg, showing gut diverticula;
i, Terminal segments of oviger with eggs.

VOLUME 95, NUMBER 2 Di)

This species is probably not rare but only rarely reported because its habitats
are seldom sampled. It was known previously only from Hawaii in the littoral
and its distribution is now extended westward to the Marshall Islands at Enewetak
Atoll.

Family Callipallenidae
Callipallene sp. cf. C. novaezealandiae (Thomson)

Pallene novae-zealandiae Thomson, 1884:246—247, pl. 14, figs. 1-4.—Hutton,
1904:247.

Callipallene brevirostris ssp. novae-zealandiae.—Stock, 1954:48—50, fig. 21a—h.

Callipallene brevirostris cf. ssp. novaezealandiae.—Arnaud, 1972:162.

Callipallene brevirostris novaezealandiae.—Child, 1975:10.

Callipallene brevirostrum ssp. novaezealandiae.—Stock, 1975a:132.

Material examined.—Large coral knoll in lagoon near Chinieero (site name
Alvin) Island in filamentous green algae at 3-5 m, 11°29’05"N, 162°23'25”E, Child
sta. 05-69, 22 Sept. 1969, 1 2 (USNM). Enewetak Island, lagoon side, in rubble
at 4m, Brock, 27 Sept. 1975, 1 d (BPBM).

Remarks.—These 2 specimens are quite close to Thomson’s species and agree
for the most part with his figures. The ovigers bear fewer denticulate spines than
either Thomson’s figure 3, or Stock’s descriptions of his specimens. The speci-
mens in hand have a denticulate spine count of 6:5:5:5, and 6:5:5:6, while Thom-
son’s female has a count of 8:8:7:8, and Stock’s male has 8:8:8:7 and his female
8:7:6:8 spines. This is the only significant difference I can find between these
Enewetak specimens and the detailed figures and descriptions given by Thomson
and Stock. It is possible that these specimens represent a new species, but with
only 2 specimens in hand and the close agreement between these and most char-
acters of Thomson’s species, I regard them provisionally as near to, if not con-
specific with his New Zealand species.

This species has never been reported from tropical western Pacific localities.
Its known distribution has been from New Zealand and Australia to the western
Indian Ocean at Madagascar and Tanzania, in mostly shallow depths. If these
specimens are in fact Thomson’s species, then the distribution is extended to the
northern hemisphere for the first time, in the Marshall Islands.

Family Nymphonidae
Nymphon micronesicum, new species
Fig. 3

Material examined.—Bikini Atoll, Namu Island, outer reef, 11°42’N, 165°17’E,
M. W. Johnson, 4 Apr. 1946, 1 2, holotype (USNM 189274).

Description.—Female (male unknown): of moderately small size, leg span al-
most 15 mm. Trunk robust, segments slightly inflated, fully segmented. Lateral
processes separated by distances slightly greater than or equal to their diameters,
only as long as their diameters, armed with few tiny setae. Neck moderately long,
with segmentation line (possible artifact) at narrowest diameter directly anterior
to oviger implantation which is slightly anterior to but not touching first lateral

278 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Nymphon micronesicum, holotype, female: a, Trunk, dorsal view; b, Ocular tubercle,
lateral view; c, Palp; d, Third leg with enlargement of ventral spine; e, Distal propodus and claws,
enlarged; f, Chela; g, Fingers of chela, enlarged; h, Oviger; i, Terminal segment of oviger with a
denticulate spine, enlarged.

processes. Ocular tubercle slightly taller than its diameter, implanted over first
lateral processes, rounded, with 2 lateral papillae and tiny apical papilla. Eyes
large, slightly pigmented. Abdomen short, erect, no taller than ocular tubercle,
armed with 2 laterodistal setae.

VOLUME 95, NUMBER 2 2H,

Proboscis a slightly swollen cylinder tapering distally to rounded lips.

Chelifore scapes large, cylindrical, 4 times as long as their diameters, armed
with several lateral setae. Chela shorter than scape, palm longer than fingers,
armed with several dorsal and distal fringe of setae. Fingers broad, immovable
finger straight, armed with 11 or 12 bicuspidate teeth. Movable finger moderately
curved, armed with 12 or 13 bicuspidate teeth. Finger tips not overlapping.

Palp short, second segment longest, terminal segment only slightly shorter.
Third segment 0.6 length of second, armed with ventrodistal setae. First and
fourth segments only slightly longer than wide. Fourth and fifth segments armed
with many setae slightly longer than segment diameter.

Oviger segments 4 and 5 subequal, armed with several distal setae. Sixth seg-
ment cylindrical, 0.5 length of fourth, armed with lateral and dorsal setae. Strigilis
4 segments armed with dorsal and lateral setae and denticulate spines in the
formula 11:9:9:10. Spines with 2 proximal large serrations and several smaller
distal serrations per side. Terminal claw as long as terminal segment, without
endal teeth or denticulations.

Leg segments slender, tibia 2 the longest segment, 0.3 longer than tibia 1 and
femur which are subequal in length. Major segments armed with short dorsal and
ventral setae increasing in numbers distally. Tibia 2 armed with 2 ventrodistal
spines, each with several endal crenulations. Tarsus and propodus short; tarsus
0.4 length of propodus, both armed with many slender setae and broader spines
on sole. Propodus without heel or larger heel spines. Major claw short, very
curved, armed with 2 rows of low conical rugosities endally. Auxiliary claws
longer than main claw, each armed with 2 rows of tiny serrations or teeth endally.

Measurements (in mm).—Trunk, first segment, 0.89; posterior 3 segments, 0.7;
width (across 2nd lateral processes), 0.67; proboscis length, 0.67; abdomen
length, 0.16; 3rd leg, coxa 1, 0.23; coxa 2, 0.64; coxa 3, 0.4; femur, 1.4; tibia 1,
1.48; tibia 2, 2.08; tarsus, 0.2; propodus, 0.47; claw, 0.1; auxiliary claws, 0.13.

Distribution.—Known only from the type-locality, Namu Island, Bikini Atoll,
in a shallow but unrecorded depth.

Etymology.—Named for Micronesia, a place of ‘small islands,’’ a number of
which form Bikini Atoll, the type-locality for this species.

Remarks.—Nymphon micronesicum is morphologically similar to N. giraffa
Loman, 1908, N. aequidigitatum Haswell, 1884, N. floridanum Hedgpeth, 1948,
and perhaps is closest to N. biformidens Stock, 1974. It shares with the latter
species a short main propodal claw with longer auxiliaries with all claws having
endal rugosities or tiny denticles, legs and palps with very similar length propor-
tions, and similar trunk and chelifore shapes. The new species differs from N.
biformidens in having a taller ocular tubercle with prominent lateral papillae, a
more cylindrical proboscis, a larger oviger claw without denticulations or rugos-
ities, longer 4th and 5th oviger segments, and more teeth on both fingers of the
chela with all chela teeth bicuspidate.

The other 3 similar species share short main claws with longer auxiliaries (those
of N. aequidigitatum lack endal rugosities), long palp terminal segments, biden-
tate chela fingers (except for N. giraffa), long second tibiae, and long oviger
terminal claws without denticulations. The new species differs from N. giraffa
in having a more compact trunk and appendages, a chela palm which is more
cylindrical than globular, a much shorter tarsus in relation to propodal length,

280 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

and very different oviger denticulate spines. Both of these species have a con-
striction or segmentation line around the narrowest part of the neck. This is
possibly an artifact due to repeated bending either during the life of the animal
or during capture.

Nymphon micronesicum differs from N. aequidigitatum in having more slender
and shorter chela fingers with many more rounded teeth, a longer tarsus in relation
to propodal length, a longer palp second segment, and a different trunk config-
uration. The new species differs from N. floridanum in having a shorter neck
without parallel sides, shorter chela fingers with fewer bicuspidate teeth, a more
cylindrical proboscis, and a terminal palp segment longer than the third segment
and almost equal to the second.

No other known Pacific species of Nymphon is as similar to this new species
as are the 4 above.

Literature Cited

Arnaud, F. 1972. Pycnogonides des récifs coralliens de Madagascar. 3. Famille des Callipallenidae.—

Téthys, Supplement 3:157-164, figures 1-6.

. 1973. Pycnogonides des récifs coralliens de Madagascar. 4. Colossendeidae, Phoxichilidiidae

et Endeidae.—Teéthys 4(4):953—960, 8 figures.

Calman, W. T. 1923. Pycnogonida of the Indian Museum.—Records of the Indian Museum 25(3):265—
299, 17 figures.

Child, C. A. 1970. Pycnogonida of the Smithsonian-Bredin Pacific Expedition, 1957.—Proceedings of
the Biological Society of Washington 83(27):287-308, 5 figures.

. 1975. Pycnogonida of Western Australia.—Smithsonian Contributions to Zoology 190:1-28,

11 figures.

. 1977. On some Pycnogonida of French Oceania.—Proceedings of the Biological Society of

Washington 90(2):440-446, 1 figure.

, and J. W. Hedgpeth. 1971. Pycnogonida of the Galapagos Islands.—Journal of Natural His-

tory 5:609-634, 8 figures.

Clark, W. C. 1963. Australian Pycnogonida.—Records of the Australian Museum 26(1):1-81, 38

figures.

. 1973. New species of Pycnogonida from New Britain and Tonga.—Pacific Science 27(1):28-

33, 3 figures.

Haswell, W. A. 1884. On the Pycnogonida of the Australian coast, with descriptions of new species.—
Proceedings of the Linnean Society of New South Wales 9:1021—1034, plates 54-57. .

Hedgpeth, J. W. 1943. Reports on the scientific results of the Atlantis expeditions to the West Indies,

under the joint auspices of the University of Havana and Harvard University. —Proceedings

of the New England Zoological Club 22:41-58, plates 8-10.

. 1948. The Pycnogonida of the western north Atlantic and the Caribbean.—Proceedings of

the United States National Museum 97 (3216):157—342, figures 4—53, 3 charts.

. 1951. Pycnogonids from Dillon Beach and vicinity, California, with descriptions of two new

species.—Wasmann Journal of Biology 9(1):105—117, 3 plates.

Hilton, W. A. 1942a. Pycnogonids from Hawaii.—Occasional Papers from the B. P. Bishop Museum,

Honolulu, Hawaii 17(3):43-55, 10 figures.

. 1942b. Pycnogonids from the Pacific. Family Ammotheidae.—Pomona Journal of EMO TIOIORY

and Zoology 34(4):93-97.

Hutton, F. W. 1904. Pp. 246-247 in Index Faunae Novae Zealandiae.—London, Dulau.

Loman, J. C. C. 1908. Die Pantopoda der Siboga-Expedition.—Siboga-Expéditie Monograph 40: fle
88, 15 plates.

Losina-Losinsky, L. 1961. [Pantopoda of the Far Eastern Seas of the U.S.S.R.}-Issledovanniya
Dalnyevostochnykh Morei SSSR, Leningrad 7:47-117, 27 figures. [In Russian].

Stock, J. H. 1953. Biological results of the Snellius Expedition XVII. Contribution to the knowledge
of the pycnogonid fauna of the East Indian Archipelago.—Temminckia 9:276—313, 18 figures.

VOLUME 95, NUMBER 2 281

. 1954. Pycnogonida from Indo-West-Pacific, Australian, and New Zealand waters.—Viden-

skabelige Meddelser fra Dansk Naturhistorisk Forening (Copenhagen) 116:1-168, 81 figures.

. 1958. The Pycnogonida of the Erythrean and of the Mediterranean coasts of Israel. Contri-

butions to the knowledge of the Red Sea. 5.—Bulletin of the Sea Fisheries Research Station,

Haifa 16:3-5.

. 1968. Pycnogonida collected by the Galathea and Anton Bruun in the Indian and Pacific

Oceans.—Videnskabelige Meddelser fra Dansk Naturhistorisk Forening (Copenhagen) 131:7-

65, 22 figures.

. 1974. Medio- and infralittoral Pycnogonida collected during the I.I.O.E. near the landbase

on Nossi-Bé, Madagascar.—Bulletin of the Zoological Museum, University of Amsterdam

4(3):11-22, 4 figures.

. 1975a. Infralittoral Pycnogonida from Tanzania.—Travais du Muséum d’ Histoire Naturelle

“Gr. Antipa,’’ Bucharest 16:127—134, 12 figures.

. 1975b. Biological Results of the University of Miami Deep-Sea Expeditions, 108. Pycnogon-

ida from the Continental Shelf, Slope, and Deep Sea of the Tropical Atlantic and east Pacific.—

Bulletin of Marine Science 24(4):957—1092, 59 figures.

Thomson, G. M. 1884. On the New Zealand Pycnogonida with descriptions of new species.—Trans-
actions and Proceedings of the New Zealand Institute 16(1883):242-—248, plates 14-16.

Department of Invertebrate Zoology (Crustacea), NHB W 323-163, Smithson-
ian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 282-291

THREE NEW SPECIES OF PYCNOGONIDA
FROM SAGAMI BAY, JAPAN!

Koichiro Nakamura and C. Allan Child

Abstract.—Three new species of Pycnogonida are described from Sagami Bay,
Japan; Ascorhynchus utinomii, Anoplodactylus shimodaensis and Anoplodacty-
lus perforatus, and their relationships to known species in the two genera are
discussed. A short resume of pycnogonid literature on Sagami Bay is given.

Introduction

Ohura or Oura Bay, locally called Nabeta Bay (34°39'30’N, 138°56’40”E), and
the immediate vicinity, is the collecting locality for 3 new pycnogonid species.
This locality is near Shimoda and is situated on the southwestern side of Sagami
Bay, an area that has received substantial coverage in pycnogonid literature.
Utinomi (1971), in his summary of known Japanese pycnogonids to that date,
listed 32 species as residents of Sagami Bay and he stated (p. 340) that ‘““Sagami
Bay and its adjacent waters is . . . the richest of all regional faunae in Japanese
waters, as represented by more than 17 genera and 46 species.’’ Fourteen of the
species listed are from off Sagami Bay proper, in deeper water.

The early literature on pycnogonids of the area includes Bohm (1879) who
described Ammothea hilgendorfi, Ascorhynchus ramipes and Propallene longi-
ceps, all from Enoshima. Ortmann (1890) described Anoplodactylus gestiens,
Ascorhynchus cryptopygium and A. bicornis (=A. auchenicum (Slater)), and list-
ed Ascorhynchus ramipes and Pycnogonum littorale var. tenue (=P. tenue (Sla-
ter)) from Sagami Bay. Ives (1891) described Ascorhynchus japonicum from Ja-
pan, listing many collecting localities, one of which was Sagami Bay. Loman
(1911) described 4 new species and listed 9 other species from Sagami Bay.

Several other authors, among them Ohshima (1936), listed subsequent records
of one or more species from this area. The next significant paper was by Hedgpeth
(1949), in his report on Japanese pycnogonids collected by the Albatross. He
listed 3 new and 3 known species from Sagami Bay (including adjacent Tokyo
Bay). Stock (1954) and Utinomi (1959, 1962, 1971) listed and described many
additional species from Sagami Bay.

Very little collecting has been done, or at least recorded in the literature,
recently for shallow water pycnogonid species in Sagami Bay, but the senior
author gathered a large collection of pycnogonids from there as a secondary result
of collecting live specimens for purposes of embryological studies. Most of the
specimens were taken from shallow water, averaging about 10 meters in depth,
in the vicinity of the Shimoda Marine Research Center, the University of Tsukuba
(formerly Shimoda Marine Biological Station, Tokyo Kyoiku University). Among
the specimens collected were the three new species of pycnogonids described in
this paper.

1 Contribution No. 378 from the Shimoda Marine Research Center, the University of Tsukuba.

VOLUME 95, NUMBER 2 283

The types are deposited in the collections of the National Museum of Natural
History, Smithsonian Institution, under the catalog numbering system of the
United States National Museum, except for several paratypes retained by the
senior author to be deposited in Japanese collections.

Pycnogonida
Family Ammotheidae Dohrn, 1881
Ascorhynchus Sars, 1877
Ascorhynchus utinomii, new species
Fig. 1

Types.—Holotype (USNM 184555), male with eggs, coll. Nabeta Bay, 7 to 15
meters, 10 July 1970. Paratypes (USNM 184556), 2 males with eggs, 3 females,
same collection as holotype. Paratypes (USNM 184557), 3 males with eggs, 1
male, 3 females, coll. Nabeta Bay, 7 to 15 meters, 13 June 1970. Paratypes
(USNM 184558), 3 males with eggs, 3 females, 1 juvenile, coll. Nabeta Bay, 7 to
15 meters, 3 October 1969. Paratypes (KN collection), 1 male with eggs, 2 fe-
males, coll. Nabeta Bay, 7 to 15 meters, 8 August 1973.

Description.—Trunk elongate, lateral processes well separated by about their
own diameter. Body smooth, without trunk tubercles, but with low rounded tu-
bercle on distal end of all lateral processes. Anterior of cephalic segment without
tubercles over chelifore insertion. Ocular tubercle placed over anterior of oviger
bases, low, rounded at tip, with 4 large eyes lightly pigmented in alcohol.

Proboscis almost half length of trunk, long and oval with one constriction
towards base giving bilobed shape, rounded in cross section.

Abdomen long, reaching almost to tips of second coxae of last pair of legs,
swollen distally.

Chelifore scape 1-segmented, longer than third palp segment, glabrous. Chela
small with atrophied tiny curved fingers.

Palp 10-segmented, basal 2 segments very small, not longer than wide, third
segment longest. Fifth segment with one large seta endally and proximally, sev-
eral shorter and 2 longer setae distally. Sixth to tenth segments densely setose
ventrally, setae subequal in length. Terminal 4 segments long, slender.

Oviger 10-segmented, with tiny terminal claw. Fourth and fifth segments long-
est, armed with row of lateral setae. Terminal 4 segments with compound spines
arranged in 2 or 3 rows, the longest row in the formula 12:9:8:10 for the holotype
and 9:8:8:10 for a juvenile.

Legs slender. First coxa with 2 dorsodistal tubercles. Coxa 2 with one low
tubercle middorsally. Femur almost as long as combined length of coxae, armed
with single dorsodistal seta. Cement glands not found. Tibia | the longest seg-
ment, armed with few tiny setae dorsally. Tibia 2 about equal in length to femur.
Tarsus slightly shorter than propodus, both without ventral setae or spines. An-
terior pair of legs with longer tarsus and propodus, armed with extremely tiny
terminal claw. Posterior 6 legs with shorter tarsus and propodus having claw
almost equal to tarsus length. Male genital pores on posterior 4 legs only. In
female, tiny pores without projections ventrally on second coxae of all legs.

Measurements.—(in mm). First trunk segment length, 1.58; total trunk length
(to tip 4th lateral processes), 3.46; trunk width (across 2nd lateral processes),
1.92; proboscis length, 1.55; abdomen length, 1.03; third leg, coxa 1, 0.53; coxa

284 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Ascorhynchus utinomii, holotype, male: a, Trunk, dorsal view; b, Trunk, lateral view; c,
Palp; d, Third leg; e, Distal segments of first leg with enlargement of tip; f, Distal chelifore, enlarged;
g, Oviger; h, Oviger terminal segments, enlarged; i, Oviger tip, further enlarged.

VOLUME 95, NUMBER 2 285

2, 0.59; coxa 3, 0.43; femur, 1.22; tibia 1, 1.78; tibia 2, 1.27; tarsus, 0.66; prop-
odus, 0.76; claw, 0.67.

Distribution.—Known only from the type-locality, Nabeta Bay in Sagami Bay,
in depths of from 7 to 15 meters.

Etymology.—This proposed new species is dedicated to the late Dr. Huzio
Utinomi, for his many contributions to our understanding of the pycnogonids of
Japan. The Japanese compound word ‘‘utinomi,’’ translated literally (pronounced
o0o-chi-noh-me), means ‘‘inside the sea.’’ This compound might in itself prove to
be a good name for a new species, but the authors prefer the patronym for this
new species.

Remarks.—This species was collected together with Propallene longiceps and
Ascorhynchus ramipes, but in fewer numbers than the others. When living, the
proboscis is light yellow in color.

This species is characterized by the shape of the proboscis, the location of the
ocular tubercle in relation to oviger implantation, lack of dorsal trunk tubercles,
and length of the abdomen. Its body size is almost the same as Ascorhynchus
minutum, but A. utinomii proves to be distinct from A. minutum in the following
details: the proboscis is bilobed with one constriction in the new species, but is
trilobed with 2 constrictions in A. minutum, and in cross section, the proboscis
is rounded in the new species and not triangular; the ocular tubercle is placed
over the anterior of the oviger implantation bulges in A. utinomii, but is entirely
anterior to the oviger bases in A. minutum; median trunk tubercles are absent in
A. utinomii, but A. minutum has small median tubercles; the abdomen is longer
in the new species, reaching almost to the tip of the second coxae of the fourth
pair of legs, but is shorter and does not reach the distal end of the first coxae in
A. minutum.

Ascorhynchus utinomii differs from the other small Japanese species such as
A. ramipes and A. auchenicum in its bilobed proboscis and lack of median trunk
tubercles. The 2 latter species have a trilobed proboscis and median trunk tu-
bercles of various sizes.

Although the cement glands of this species are not evident, they are probably
a row of extremely tiny pores such as those found on some of the larger species
of the genus.

Family Phoxichilidiidae Sars, 1891
Anoplodactylus Wilson, 1878
Anoplodactylus shimodaensis, new species
Fig. 2

Types.—Holotype (USNM 184559), male with eggs, coll. Nabeta Bay, 7 to 15
meters, 13 May 1970. Paratypes (USNM 184560), 2 males with eggs, 3 females,
same collection as holotype. Paratype (USNM 184561), 1 male with eggs, coll.
Nabeta Bay, 7 to 15 meters, 3 October 1969. Paratypes (USNM 184562), 1 male
with eggs, 5 females, coll. off Kisami, 30 meters, 22 May 1970. Paratypes (USNM
184563), 3 males, 1 female, coll. Nabeta Bay, 7 to 15 meters, 19 January 1971.
Paratypes (KN collection), 3 males with eggs, 2 females, coll. Nabeta Bay, 7 to
15 meters, 5 October 1970.

Description.—Trunk robust, first and second segmentation lines complete. No

286 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Anoplodactylus shimodaensis, holotype, male: a, Trunk, dorsal view; b, Trunk, lateral
view; c, Third leg, with enlargement of cement gland and propodal heel; d, Chela.

VOLUME 95, NUMBER 2 287

segmentation between third and fourth trunk segments. Lateral processes long,
over twice as long as their diameters, separated by their diameters or less, without
tubercles, armed with 3 or 4 dorsodistal setae. Fourth pair of lateral processes
shortest. Neck narrow, ocular tubercle about 1.5 times as tall as width at base,
rounded at tip, with low apical cone and 4 large lightly pigmented eyes. Lateral
sensory papillae large, prominent.

Proboscis short, robust, blunt at tip with constriction near distal fourth of
length.

Abdomen short, erect, approximately as tall as ocular tubercle, tapering to
rounded tip, armed with several short setae.

Chelifore scape straight proximally, strongly curved ventrodistally, armed with
few setae. Chela small, with curved fingers overlapping at tips. Movable finger
with 2 tiny tooth-like bumps, immovable finger without teeth.

Oviger of 6-segments, third with proximal construction. Terminal 2 segments
armed with many setae shorter than segment diameter.

Legs moderately long. First coxa with distal setae, second coxa longest. Femur
longest of major segments, armed with few setae and a long dorsodistal seta.
Cement gland a stout tube shorter than segment diameter, opening dorsally at
distal fourth of segment. First and second tibiae almost equal in length, armed
with single long dorsodistal seta each. Tarsus short, almost triangular, armed
with several ventrodistal setae. Propodus slightly curved, sole straight, with
marked heel armed with 2 short stout spines and 2 setae. Sole with propodal
lamina over entire length flanked by few short setae. Claw 0.7 propodal length,
slender. Auxiliaries tiny.

Measurements.—(in mm). Trunk length (without chelifores), 1.45; trunk width
(across 2nd lateral processes), 1.26; proboscis length, 0.59; abdomen length, 0.24;
third leg, coxa 1, 0.35; coxa 2, 0.53; coxa 3, 0.44; femur, 0.91; tibia 1, 0.82; tibia
2, 0.79; tarsus, 0.11; propodus, 0.5; claw, 0.36.

Distribution.—Known from the type-locality, Nabeta Bay on Sagami Bay in
depths of from 7 to 15 meters. Also found off Kisami in 30 meters.

Etymology.—Named for the town of Shimoda, near where the species was
collected. The town of Shimoda was made famous, if fleetingly, as the first place
where Commodore Perry landed in 1853, to open diplomatic relations with Japan.

Remarks.—This species is easily distinguishable among Japanese species of
Anoplodactylus by its small robust size and shape, the short barrel-shaped pro-
boscis with construction, and the wide distally-placed tube of the cement gland
in the male. No other known Japanese species in this genus has such robust
lateral processes separated from each other by their own diameter, in combination
with a short stout proboscis. The single femoral cement glands of this species are
placed in a relatively rare distal position. Where the cement gland configuration
is known, there are few species in this crowded genus which have the cement
gland placed in the distal fourth or even third of the femur. We are unable to find
any other known species of the genus from Japan or vicinity with a broad tubular
cement gland placed this far forward on the femur.

There are several known species of Anoplodactylus with similarities shared by
this species, but none of them have the above characters in combination with
chela fingers without teeth, no genital spurs on the second coxae of the third and
fourth legs, and a moderate length ocular tubercle with conical apex.

288 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Anoplodactylus perforatus, holotype, male: a, Trunk, dorsal view; b, Trunk, lateral view;
c, Terminal segments of oviger, enlarged; d, Third leg, 3 cement glands enlarged; e, Third leg terminal
segments; f, Propodal lamina enlarged; g, Chela.

VOLUME 95, NUMBER 2 289

Anoplodactylus perforatus, new species
Fig. 3

Types.—Holotype (USNM 184564), male with eggs, coll. Nabeta Bay, 7 to 15
meters, 3 June 1970. Paratypes (USNM 184565), 1 male with eggs, 2 males, 2
females, 1 juvenile male, same collection as holotype. Paratype (KN collection),
1 male with eggs, coll. Nabeta Bay, 7 to 15 meters, 3 October 1969. Paratype
(USNM 184566), 1 male, R/V Tansei Maru, cruise KT 69-12, sta. 21, 35°00'54’N,
139°08'36’E, 113 meters, 17 July 1969.

Description.—Trunk elongate, fully segmented between first 3 segments, third
and fourth unsegmented. Lateral processes separated by less than their diameters,
armed with 2 or 3 dorsodistal setae, without tubercles. Neck short, slender.
Ocular tubercle rounded with conical apex, 1.5 times taller than width at base,
with 4 large darkly pigmented eyes.

Proboscis slender, blunt at tip, slightly constricted distally.

Abdomen erect, tapering to rounded tip, glabrous, about equal in length to
ocular tubercle.

Chelifore scape slender, slightly curved laterally, armed with several dorsal
setae. Chela large, palm cylindrical, with 2 or 3 setae. Fingers placed almost at
right angles to palm, curved, overlapping at tips, armed laterally and dorsally
with several setae, without teeth.

Oviger 6-segmented, long, third segment longest, with proximal constriction.
Distal 2 segments armed with many curved setae. Fifth segment armed with single
short recurved spine endally.

Legs slender. Coxae armed with several dorsal and ventrodistal setae. Second
coxae without genital spur. Femur almost equal in length to tibia 1, both with
small dorsodistal tubercle bearing a seta. Femoral cement gland openings dorsal,
arranged in single row of cribellate pores numbering 17 to 25 per leg. First tibia
slightly longer than second, both armed with many dorsal, lateral and ventral
setae shorter than segment diameter and single dorsodistal seta longer than seg-
ment diameter. Tarsus short, semitriangular, with several ventral and single dor-
sal setae. Propodus moderately curved, with marked heel bearing 2 spines. Sole
without spines proximally, with about 10 spines and several lateral setae along
distal three-quarters of length and very short propodal lamina distally, armed
dorsally and laterally with several setae. Terminal claw long, moderately curved,
about two-thirds propodal length. Auxiliary claws tiny.

Measurements.—(Gn mm). Trunk length without chelifores, 1.97; trunk width
(across 2nd lateral processes), 1.27; proboscis length, 1.34; abdomen length, 0.37;
third leg, coxa 1, 0.47; coxa 2, 0.9; coxa 3, 0.65; femur, 1.68; tibia 1, 1.64; tibia
2, 1.4; tarsus, 0.13; propodus, 0.78; claw, 0.54.

Distribution.—Known from the type-locality, Nabeta Bay in from 7 to 15 me-
ters, and from off Shimoda in 113 meters.

Etymology.—Named for the perforated appearance of the 17 to 25 cement
glands along the dorsal surface of the femur.

Remarks.—This species closely resembles Anoplodactylus gestiens, except
that it is not quite as slender or tenuous and has many cement glands instead of
the single gland. A set of figures of A. gestiens is included here (Fig. 4) because
this species has never, in our opinion, been adequately figured in the literature.

290 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Anoplodactylus gestiens, specimen from TS-69, station 21, male: a, Trunk, dorsal view;
b, Trunk, lateral view; c, Third leg; d, Terminal segments of third leg; e, Chela.

There is no known species of Anoplodactylus having so many cement glands
per femur. The nearest species in number of cement glands is A. cribellatus which
is reported in the literature to have fifteen or so per leg. All other known species
have 10 or less glands per femur. The 14 known species of Anoplodactylus with
multiple glands are as follows: A. cribellatus Carpenter, with about 15 glands; A.
longiformis Child, with 8 to 10; A. australis (Hodgson), about 7; A. xenus Stock,
about 8; A. pycnosoma (Helfer), from 5 to 8; A. multiclavus Child, 3 to 5; A.
glandulifer Stock, 2 to 4; A. angulatus (Dohrn), 4 or 5; A. dentimanus Stock, 3;
A. oculatus Carpenter, 5; A. virescens (Hodge), 2 or 3; A. robustus (Dohrn), 3;
A. longiceps Stock, 2; and the present new species with from 17 to 25 glands.

VOLUME 95, NUMBER 2 291

There are several species of the closely related genus Phoxichilidium which have
multiple cement glands on the legs of males.

The proposed new species has no other conspicuous characters to set it apart
from many of the genus, but the cement gland number and shape have proved to
be sufficiently stable in providing a primary source of differentiation among the
many species of Anoplodactylus.

Acknowledgments

We acknowledge the assistance and generosity of the Shimoda Laboratory
staff, particularly Hajime Ueda, without whose help this collection of new species
could not have been made. We are grateful also to Dr. Koichi Sekiguchi of
the University of Tsukuba, and chief scientist of the Tansei Maru Expedition,
for his collaboration in collecting material.

Literature Cited

Bohm, R. 1879. Uber zwei neue von Herrn Dr. Hilgendorf in Japan gesammelte Pycnogoniden.—
Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin 1879(4):53-60, figs.
a—c.

Hedgpeth, J. W. 1949. Report on the Pycnogonida collected by the Albatross in Japanese waters in
1900 and 1906.—Proceedings of the United States National Museum 98(3231):233-321, figs.
IS=51,

Ives, J. E. 1891. Echinoderms and arthropods from Japan.—Proceedings of the Academy of Natural
Sciences of Philadelphia, 1891—1892:210-221, plate XII.

Loman, J. C. C. 1911. Japanische Podosomata; Beitrage zur Naturgeschichte Ostasiens, herausgeben
von F. Doflein.—Abhandlungen der Kaiserlich Bayerischen Akademie der Wissenschaften
(Mathematisch-Naturwissenschaftliche Klasse), Suppl. 2(4):1—18, pls. I-II.

Ohshima, H. 1936. [A list of Pycnogonida recorded from Japanese and adjacent waters.]|—Dobut-
sugaku Zasshi 48(8—10):861-869. [In Japanese.]

Ortmann, A. E. 1890. Bericht tiber die von Herrn Dr. Déderlein in Japan gesammelten Pycnogoni-
den.—Zoologische Jahrbiicher (Systematik) 5(1):157-168, pl. XXIV.

Stock, J. H. 1954. Pycnogonida from Indo-West Pacific, Australian and New Zealand waters. Papers
from Dr. Th. Mortensen’s Pacific Expedition 1914—-1916.—Videnskabelige Meddelelser fra
Dansk Naturhistorisk Forening i Kjobenhavn 116:1-168, figs. 1-81.

Utinomi, H. 1959. Pycnogonida of Sagami Bay. Publications of the Seto Marine Biological Laboratory

7(2):197—222, figs. 1-9.

. 1962. Pycnogonida of Sagami Bay—Supplement.—Publications of the Seto Marine Biological

Laboratory 10(1):91-104, figs. 1-6.

. 1971. Records of Pycnogonida from shallow waters of Japan.—Publications of the Seto

Marine Biological Laboratory 18(5):317—347.

(KN) 2-25-3, Nakamachi, Musashino-shi, Tokyo 180, Japan; (CAC) Depart-
ment of Invertebrate Zoology (Crustacea), NHB W323, stop 163, Smithsonian
Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 292-296

A GYNANDROMORPH OF THE JAPANESE PYCNOGONID
ANOPLODACTYLUS GESTIENS (ORTMANN)

C. Allan Child and Koichiro Nakamura

Abstract.—A single sexual mosaic or gynandromorph specimen was found
among a very large series of the Japanese pycnogonid Anoplodactylus gestiens
(Ortmann). The rarity and morphology of the specimen are examined and de-
scribed, and its relationship with other arthropod and pycnogonid gynandro-
morphs is discussed.

Pycnogonids are normally dioecious and sexually dimorphic with testes or ova-
ries situated in each leg. In the few instances where sexual habits have been
observed, the male accepts eggs released from the female’s sexual pores, fertilizes
them externally, cements the eggs together by use of femoral cement glands, and
carries the eggs cemented to his ovigers until after hatching. The sexual habits,
as far as we have been able to discover in the literature, are unknown for many
genera, including those few species having parasitic or commensal larvae and
young not carried by the male. It is therefore significant to discover a single
apparent gynandromorph specimen among a very large sample of the pycnogonid
Anoplodactylus gestiens (Ortmann, 1891), collected in Sagami Bay, Japan.

The occurrence of hermaphrodites and gynandromorphs among the Pycnogon-
ida is extremely rare according to the literature. There are only three examples
of these abnormalities on record. The first two references (to one hermaphroditic
species and a gynandromorphic specimen of another species) are reviewed by
Child (1978:134), and the third example, a gynandromorph series, is discussed in
the same paper (Child 1978:135-141, figs. 1-4). Such abnormalities are apparently
equally rare in other marine arthropods and there is only a very limited literature
on the subject (Chace and Moore 1959:226—231, figs. 1-4; Froglia and Manning
1978:700, fig. 5; Manning and Holthuis 1981:62, fig. 14). The majority of papers
concern bilateral gynandromorphs where half the animal has male characters
including obvious coloration, and the other half has female characters with female
coloration. It is to be assumed that where coloration is not a factor and where
micro- or semimicroorganisms are concerned, a small number of hermaphrodites
or gynandromorphs would pass unobserved, particularly when examining a large
series of these organisms. It was therefore rewarding when one of us (Nakamura)
discovered a single abnormal specimen while determining the sex of a series of
598 specimens of Anoplodactylus gestiens (Ortmann).

This pycnogonid is a fairly commonly captured endemic in the shallow waters
of Japan and is listed in most taxonomic reports concerning Japanese collections.
The collection in which this single gynandromorph was found was taken in a
depth of 113 meters during a dredging-trawling survey by the R/V Tansel Maru
of the Ocean Research Institute of the University of Tokyo, while sampling for
certain common pycnogonids to be used in seasonal fluctuation studies of both
numbers and sex frequency over a 28 month period (Nakamura and Sekiguchi,
in press).

VOLUME 95, NUMBER 2 293

Fig. 1. Anoplodactylus gestiens, gynandromorph appendages: A, Normal female leg with ova; B,
Leg L 4 of gynandromorph with enlargement of cement gland; C, Normal male leg; D, Proximal
segments of gynandromorph ovigers, left at top, right at bottom. Abbreviations: ov, ovary with ova;
id, intestinal diverticulum; cg, cement gland and tube; t, testes; sp, sexual pore.

Description of the Specimen

This specimen has characters of both male and female pycnogonids of the genus
Anoplodactylus. Normally, gut diverticula extend out through most segments of
each leg of a pycnogonid. The ovaries of the female lie dorsally to these diver-
ticula and are usually confined to the first four segments of each leg (Fig. 1A). In

294 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

the male, the testes lie ventrally to the diverticula and are usually confined to the
three coxae or the second and third coxae only (Fig. 1C). Testes are normally
found in the posterior four legs or the posterior pair of legs only. Both sexes have
sexual discharge pores placed ventrally on the second coxae; the female’s on all
eight legs and the male’s corresponding to the occurrence of testes in the posterior
legs. Normally only males of this genus have ovigers, the egg-carrying appendages
to which the extruded eggs are cemented (except for a few enigmatic females of
Anoplodactylus jonesi Child, 1974).

This apparent gynandromorph has ovigers like the male but with the left one
slightly deformed (Fig. 1D), has female ovaries distended with ova in all eight
legs, has sex pores on all eight legs, and has a femoral cement gland present but
reduced or atrophied on leg L 4 (Fig. 1B).

Each set of coxae was carefully examined for the presence of testes, but aside
from the darker opaque tubes of the gut diverticula and ovaries, none of the eight
legs have any matter resembling a testes in the coxae. Unlike the hermaphrodite
species (of a different genus) found by Marcus (1952), this specimen is apparently
incapable of self-fertilization and thus is more properly termed a sexual mosaic
or gynandromorph rather than an hermaphrodite.

The right oviger is normal to all appearances, but the left is slightly shorter
with the two proximal segments reduced in diameter. The second segment tapers
proximally and is expanded distally, forming a club-like shape. The normal ce-
ment gland in Anoplodactylus gestiens is an oblong opaque sac placed mid-dor-
sally in the femur and extending outside the integument in the form of a slanted
tube as long as one-third the diameter of the segment. The cement gland present
on leg L 4, the single leg out of eight with the gland in this specimen, is a tiny
tube expanded at its base, and measures in length less than one-fourth the femur
diameter. The gland within the segment has the shape of a small dark bifurcated
tube. The space which would normally be taken up by a full-sized gland is lighter
colored than the surrounding tissues as though the space were empty. The smaller
cement gland is placed more toward the distal end of the femur than the normal
mid-dorsal placement in the male leg. The seven other legs have no suggestion
of cement glands and are apparently normal in all respects for an ovigerous
female. All eight legs have normal sexual discharge pores on the second coxae.

There are other differences between this specimen and our series of Anoplo-
dactylus gestiens specimens. These differences in the gynandromorph appear to
fall within the range of non-sexual variation in the species. This is verified, except
for a difference in specimen size, by the large series in hand. The ocular tubercle
is shorter in the gynandromorph, but this tubercle varies in length through the
series. The propodal lamina is slightly longer than usual in this specimen and
corresponds to Stock’s (1954:69) figure 3la of a propodus from a young female
of the species. Some of the female lamina are shorter than this in our series of
specimens.

The single major difference in the gynandromorph specimen is its reduced size.
It measures only about three-fourths the size of a normal male of the species,
and in pycnogonids the male is almost always smaller than the female. This could
be attributed to the mosaic characters of the specimen, but with other sexual
mosaic pycnogonids (Child 1978), little or no difference in size was found between
normal and abnormal specimens.

VOLUME 95, NUMBER 2 295

Discussion

This single gynandromorph among 598 specimens of the same Anoplodactylus
species affords no hint of the cause or origin of this abnormality in the population
as sampled. Records were not made at the time of capture concerning associated
fauna or bottom conditions, but the depth of 113 meters at which the sample was
taken would preclude the influence of wave action, ship’s wake, or other surface
phenomena as a disturbance in the embryonic determination of sexual characters.
On the other hand, it is entirely possible that this specimen developed in a dif-
ferent habitat and depth and only reached the sampling depth as an adult.

The fact that this isolated abnormality is not unique among the pycnogonids is
of some interest. Gynandromorphy has been discovered among species of the
genera Anoplodactylus and Ascorhynchus. Anoplodactylus gestiens is the second
species of this genus in which gynandromorphs are known, the other being A.
portus Calman, 1927 (Child 1978). The littoral gynandromorphs of A. portus were
possibly caused by agitation of the eggs during the time of early embryonic de-
termination and development by fast ships’ propellers. This could account for the
random nature of visible abnormalities rather than a bilateral appearance of the
resulting gynandromorphs. A random gynandromorph pycnogonid has one leg
with male characters, the next with female characters, the next with both male
and female characters, and the next with only female characters, and so on, with
no apparent pattern. This gynandromorph of A. gestiens has only one random
character, that of the reduced cement gland on the left fourth leg. All of the legs
have developing eggs and seven of the eight legs are normal female legs in ap-
pearance, while the specimen has fully developed male ovigers with the left oviger
slightly deformed. There is no randomness in these last characters even though
the presence of both ovigers and ovaries in a single specimen is contrary to the
definition of this genus. The fact that both the reduced cement gland and the
malformed oviger occur on the left side of the specimen may be due to coinci-
dence or it may signify a form of bilateral mosaic, but there is no further evidence
to support bilateralism.

The two species with abnormalities in the genus Ascorhynchus are A. abyssi
Sars, 1877, and A. corderoi du Bois-Reymond Marcus, 1952. Losina-Losinsky
(1964) described but did not figure an apparent bilateral of A. abyssi having male
legs on one side and female legs with eggs on the other. Marcus, in describing
her new species A. corderoi, listed several specimens with both testes and ovaries
in the same legs, making this the only true hermaphrodite species known.

There is one other species of Anoplodactylus, A. jonesi, in which a few spec-
imens out of all of those known, while not apparently gynandromorphs, have
female legs lacking cement glands while at the same time having male ovigers.
Based on our knowledge of the genus, there is as little explanation for this sexual
combination as there is for the gynandromorph in A. gestiens. Perhaps it will be
necessary to redefine the genus based on this occasional presence of otherwise
absent characters for a few of the known species.

Acknowledgments

We express our gratitude to Dr. F. A. Chace, Jr., Department of Invertebrate
Zoology, Smithsonian Institution, for his critical review of the manuscript, to

296 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Professor Koichi Sekiguchi of the University of Tsukuba and chief scientist of
the Tansei Maru Expedition, for his collaboration in collecting material (with
KN), to Mr. H. Ueda of the M.R.C. for his excellent technical assistance in
collecting the specimens (with KN), and to the Department of Invertebrate Zoology
(Crustacea), Smithsonian Institution, for use of visitor facilities (by KN).

Literature Cited

Chace, F. A., Jr., and G. M. Moore. 1959. A bicolored gynandromorph of the lobster, Homarus
americanus.—Biological Bulletin 116(2):226—231, figs. 1-4.

Child, C. A. 1978. Gynandromorphs of the pycnogonid Anoplodactylus portus.—Zoological Journal
of the Linnean Society 63:133-144, 4 figures.

Froglia, C., and R. B. Manning. 1978. Brachynotus gemmellari (Rizza, 1839), the third Mediterranean
species of the genus.—Proceedings of the Biological Society of Washington 91(3):691-705, figs.
lS),

Losina-Losinsky, L. 1964. [Pantopoda from the collections of the F. Litke in 1955 and the Ob in
1956. Scientific results of the oceanographic expeditions to the northern parts of the Greenland
Sea and neighboring regions of the Arctic Basin in the years 1955—1958].—Trudy arkticheski
i antarkicheski Nauchnovo-issiedovatetskovo institut 259:330—339 [In Russian].

Manning, R. B., and L. B. Holthuis. 1981. West African Brachyuran Crabs (Crustacea: Decapoda).—
Smithsonian Contributions to Zoology 306:1—xii, 1-379, figs. 1-88.

Marcus, E. du B.-R. 1952. A hermaphrodite pantopod.—Anais da Academia Brasileira de Ciéncias
24(1):23-30.

Nakamura, K., and K. Sekiguchi. (in press). Seasonal occurrence of four species of pycnogonids in
Nabeta Bay, Shimoda, Japan.

Ortmann, A. 1891. Bericht tiber die von Herrn Dr. Doderlein in Japan gesammelten Pycnogoniden.—
Zoologische Jahrbticher (Systematik) 5(1):157-168, plate 24.

Stock, J. H. 1954. Pycnogonida from the Indo-West-Pacific, Australian and New Zealand waters.
Papers from Dr. Th. Mortensen’s Pacific Expedition 1914—1916.—Videnskabelige Meddelelser
fra Dansk Naturhistorisk Forening i Kjgbenhavn 116:1-168, 81 figures.

(CAC) Department of Invertebrate Zoology, Smithsonian Institution, Wash-
ington, D.C. 20560; (KN) 2-25-3 Nakamachi, Musashino-shi, Tokyo 180, Japan.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 297-318

THE ENTOCYTHERID OSTRACOD FAUNA OF
NORTHERN GEORGIA

Horton H. Hobbs, Jr. and Daniel J. Peters

Abstract.—The entocytherid ostracod fauna of the Tennessee and Coosa river
basins (exclusive of the Tallapoosa sub-basin of the latter) in Georgia consists of
13 species that infest one or more of 26 of the 27 epigean crayfishes frequenting
these watersheds. None of the ostracods is restricted to the area, and there is no
evidence that any of them require a specific host. Spot maps depicting the locality
records for each ostracod, summaries of their ranges, and their associations with
the crayfish hosts and other entocytherids occurring in the area are tabulated. A
key for recognizing the ostracods is also included.

Introduction

During a survey of the crayfish fauna of Georgia (Hobbs 1981), many of the
entocytherid ostracods infesting these decapods were preserved for future study.
Those available from the Coosa (exclusive of the Tallapoosa) and Tennessee river
basins, an area herein designated as “‘northern Georgia,’’ constitute the material
on which this is based.

The entocytherid ostracods are obligate symbionts of aquatic amphipods, iso-
pods, crayfishes, and crabs. Only members of the nominate subfamily occur in
Georgia, and all are associated with crayfishes. The entocytherid fauna of the
area treated here consists of 13 species that infest one or more of 26 of the 27
epigean species of cambarine crayfishes frequenting the Coosa and Tennessee
watersheds.

Not one of the 13 entocytherids is restricted to northern Georgia, but we an-
ticipate that the southeastern limits of the range of at least two of them (Asce-
tocythere bouchardi and Dactylocythere prominula) will prove to lie within the
area. Few of the members of the genus Dactylocythere are expected to be found
to the south except in the headwaters of the Chattahoochee and Savannah rivers.

The area under consideration.—The Coosa and Tennessee rivers (Fig. 1) drain
some 12,000 and 7250 square kilometers, respectively, of the northern part of
Georgia. Within this area, there are segments of four of the five physiographic
regions represented in the state (Fig. 2). Furthermore, elevations range from
approximately 152 to 1450 meters. |

The headwater streams and most of the creeks and rivers outside of the Ridge
and Valley Province flow, for the most part, over granitic or sandstone beds and
thus have low mineral content. Those coursing at lower elevations of the Ridge
and Valley Province flow over eroded beds of Paleozoic limestone, dolomite, and
shale and are enriched by calcium and magnesium salts.

A broad array of habitats ranging from seepage areas and mountain rills to
rather large, sluggish streams and crayfish burrows is available to the ento-
cytherids in this section of the state, and whereas all appear to have been ex-
ploited by the ostracods, we have been able to recognize few correlations of their

298 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Little Tennessee

ae

AN

Coosawattee SS.
ne NGA ud ‘
stanaula

es

3 “ |
JC Chattahoochee ane

==

se

Fig. 1. River basins in northern Georgia.

distribution with ecological conditions. Unlike entocytherid faunas in other areas
in which unique species are symbionts of those crayfishes largely restricted to
burrows, the ostracods infesting the burrowing crayfishes in northern Georgia are
members of some of the same species that are found on the stream-dwelling
crayfishes frequenting nearby epigean habitats. The ostracods occurring in the
physiographic provinces and drainage basins are summarized in Table 1.

Entocytherid associates and infestations.—As we stated in our summary of the
entocytherids of North Carolina (Hobbs and Peters 1977:4), ‘“‘In . . . [northern
Georgia], as elsewhere, frequently more than one entocytherid infests the same
crayfish, and in this study the ostracods were, for the most part, obtained from
collections of crayfishes in which all of the specimens collected in one locality
were preserved in the same container. Thus, if two or three species of crayfishes
were obtained at one station, we were unable to determine which of the ostracods
retrieved from the container infested which host species. The only instances in
which we can be certain that an entocytherid was associated with a particular
host species are those in which collections contain only one species of crayfish.
In Table 2, the associations indicated are based in part upon records of ostracods
occurring in the same locality, perhaps infesting more than one species of crayfish
(open circles) rather than occurring on the same host species (solid circles).’’ Our
collections were obtained from 154 localities (Fig. 3).

Whereas an obligate association with one or more crayfishes exists for all of
the entocytherids of the area, evidence exists that none requires a specific host.

VOLUME 95, NUMBER 2 299

85° 84°

OO. se
= ORR T, SS Sy >
Eee ys
“or a
| St

riA—--

~
7 J
(4
| NS
we
i eX S

| \
We IN
; DaN
Wty, Oe Ae She 7c h Yy
i 6 Piedmont ie y
, ; J l ; sere a
ee i Ps 2 eK

AA
4,

SS /

34°

y 7 _

Fig. 2. Physiographic provinces in northern Georgia.

The only ostracod known to be associated with a single host species in the region
is Dactylocythere prominula which was found in a single locality on a small
mountain stream in which only one crayfish, Cambarus (C.) bartonii, was col-
lected. In several localities in Tennessee, however, it has been found on other
crayfishes (Hobbs and Walton 1977:609), and we believe it highly likely that it
will be found on species other than C. (C.) bartonii in northern Georgia.

Entocytherid distribution in northern Georgia.—The numbers of entocytherids
(Table 1) and crayfishes (Hobbs 1981:table 2) occurring in the four provinces in
northern Georgia are as follows:

Provinces Entocytherids Crayfishes
Appalachian Plateau 6 8
Ridge and Valley 10 19
Blue Ridge 7 13
Piedmont i 8

Obviously the Ridge and Valley Province supports the largest number of both
entocytherids and crayfishes: 10 of the 13 known ostracods and 20 of the 27
crayfishes. Despite the relatively small area of northern Georgia encompassed by
the Appalachian Plateau, almost half of the species comprising the entocytherid
fauna have been found within it.

Entocythere elliptica and Uncinocythere simondsi are the most widespread
entocytherids in northern Georgia, both occurring in all of the physiographic

300 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Distribution of entocytherids in the physiographic provinces and drainage basins of north-

ern Georgia.
ENTOCYTHERIDS

elliptica
illinoisensis
simondsi

nae

7a)

~

=

23 S

DW

as 3

—D =~

zy 3

aS >

S S
iS

O 3S

aS ~

= Q

3
~~
3
©
La}
3
el
+S
Q

&
is)
=
a
3
ie)
~
o
—
+
QA

3

2/8
B3/ 8
Ss
of
o|e
£| 2
+5] 45
=)

Ss
~
D

~
=~)
iva)
+
=

provinces _| | | | | | | | | || | |
lel OBO

E
E
U

| An.telmoecea |
As. bouchardi
C. cyma

Appalachian Plateau| |@| | | jel | je

Ridge and Valley _—|@| |@jejejeie/ |e |elele
Blue Ridge | | |e} jeje! |e! jeje ie
e| |e e| |e

Piedmont

DRAINAGE BASINS
Tennessee

C)
e@
TELE
Om
Hiwassee Be

i
e@|
el eel
Little Tennessee esl
Coosa eae :
| | jo] jeje) jele| |e
bleed
Om
iol ad

Coosawattee
Etowah
Chattooga

Conasauga
| | jeje! | | jeje |e
OBOE BORO

provinces and in all major drainage basins except the Little Tennessee. Only
slightly less broadly distributed is Dactylocythere falcata, which has not been
found on the Appalachian Plateau. In contrast, Ascetocythere bouchardi, Dac-
tylocythere brachystrix, Dt. prominula, Dt. suteri, and Entocythere illinoisensis
have the most restricted ranges. The first of these has been found in four localities
in the Tennessee Basin lying on the Appalachian Plateau. Dactylocythere bra-
chystrix is a little more widespread, for while known only from the Tennessee
Basin, it occurs in streams of both the Appalachian Plateau and Ridge and Valley
provinces. Dactylocythere prominula was discovered in only a single locality,
that lying in the Conasauga Basin of the Blue Ridge Province. Dactylocythere
suteri was obtained in several localities in the same river basin, but in the Ridge
and Valley instead of the Blue Ridge Province. Like Dt. prominula, E. illinois-
ensis was found in a single locality in the area, but one lying in the Tennessee
watershed of the Ridge and Valley Province.

VOLUME 95, NUMBER 2

301

Table 2.—The entocytherids and their crayfish hosts in northern Georgia (solid circles = known
host-commensal relationships; open circles = occurrence in same locality; see ‘‘Entocytherid asso-

ciates and infestations’’ for more detailed explanation).

HOSTS

C. (D.) latimanus

ENTOCYTHERID

e@[O[O[Ole[e[olo] Dr. falcata

QISSes Sc SCRSeBPERBS ORONO SC ER4a minma
je; | | | | feo | fe} | Tt | fe@fol TT folo} je} je} dt. teptophytax

SJEbSS 228 SREP ROCOSS RASA BE Aenean
Covet LEceese hE SS |_Ce TEs
Sa A538 SESEEREC CESSES RA RAR RRS
| jojo! jo} | fof fol | fol | Tt | | fojet | [ojo] le] Dt. mecoscapha __

ISSR SRS ECEREESER Pea sams
SSE SS ST EROOSEE SENN aOR RE aes: |
|| | | jo} fefofe] | | fefojofelo] | | jejol jo | le] Dn. donnaldsonensi

e@/olo| jelelele| jojejoje! ol [olelcle[clejolejelcle| £. elliptica
JS sacs tee cokes peewee.

Lllinoisensis

ojojoy je@jo} jo} jojejojojo] | [ejejoloje/oj[ojejejolel U. simonds:

302 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Localities from which entocytherids have been examined.

Finding Ankylocythere telmocea in the Tennessee Basin, and especially in the
Ridge and Valley Province, came as a real surprise to us for elsewhere in its
range it has been reported only from the Piedmont and Coastal Plain provinces
(see Hart and Hart 1974:32-33). This isolated locality, and perhaps others in the
province, should be investigated further to assure the validity of the occurrence
of the species in the Tennessee River basin.

In respect to their hosts, Entocythere elliptica is apparently the least selective
of the entocytherids occurring in northern Georgia, having been definitely asso-
ciated with 13, and possibly occurring on as many as 23, of the 27 crayfishes
inhabiting the area. Only slightly less selective are Dactylocythere falcata and
Uncinocythere simondsi. The former has been found on 12 species of crayfishes
frequenting the area, and the possibility exists that 10 additional host species are
also infested by Dt. falcata. Uncinocythere simondsi has found the exoskeleton
of all of the crayfishes acceptable for colonization except perhaps those of Cam-
barus (J.) nodosus, C. (J.) distans, C. (P.) georgiae, C. (P.) parrishi, and O.
forceps.

Although most studies have indicated that crayfishes of the genus Orconectes,
in general, support smaller numbers of ostracods than do most members of other
cambarine genera, we are not confident that Orconectes forceps lacks ento-
cytherid infestations. The few collections of this crayfish from Georgia streams
that are available to us were not maintained in the original containers in which
they were collected, and the fluid in which they were killed had been discarded.

VOLUME 95, NUMBER 2 303

Table 3.—Associations of entocytherids in northern Georgia (solid circles = known utilization of
same host species in at least one locality; open circles = occurrence in same locality; see ‘‘Ento-
cytherid associates and infestations’’ for more detailed explanation).

- ENTOCYTHERID
ASSOCIATES

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Cambarus (J.) distans has been found in only two localities, both on an inter-
mittent stream in the northern part of Dade County, and in one of them it is
infested with Donnaldsoncythere donnaldsonensis, one of the most widely rang-
ing entocytherids in the eastern half of the United States. Cambarus (P.) geor-
giae, known from a single locality in the Georgia segment of the Little Tennessee
River, harbors two of the three ostracods known to occur in the headwaters of
that stream.

The burrowing Cambarus (D.) cymatilis and C. (J.) nodosus and the riffle-
inhabiting C. (H.) manningi are infested by three, two, and three entocytherid
Species, respectively, but the ostracods on them are among the commonest in the
area, infesting a comparatively large variety of crayfish hosts (Table 1).

In contrast, the least ecologically restricted crayfish in the area, Cambarus (D.)
striatus, has been found to host five and possibly nine entocytherids. Cambarus
(C.) bartonii and C. (P.) coosae harbor six and five species of entocytherids,
respectively, and possibly an additional one occurs on each.

304 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

The host species in the area have been treated by Hobbs (1981), and synony-
mies, ranges, and notes on their biology are recorded therein. The epigean cray-
fish fauna of northern Georgia, as delimited here, consists of the following
species: Cambarus (Cambarus) bartonii (Fabricius, 1798); C. (Depressicamba-
rus) cymatilis Hobbs, 1970; C. (D.) latimanus (LeConte, 1856); C. (D.) striatus
Hay, 1902; C. (Hiaticambarus) coosawattae Hobbs, 1981; C. (H.) fasciatus
Hobbs, 1981; C. (H.) girardianus Faxon, 1884; C. (H.) longirostris Faxon, 1885;
C. (H.) manningi Hobbs, 1981; C. (H.) speciosus Hobbs, 1981; C. (Jugicam-
barus) conasaugaensis Hobbs and Hobbs, 1962; C. (J.) distans Rhoades, 1944;
C. (J.) nodosus Bouchard and Hobbs, 1976; C. (J.) parvoculus Hobbs and Shoup,
1947; C. VJ.) unestami Hobbs and Hall, 1969; C. (Lacunicambarus) acanthura
Hobbs, 1981; C. (Puncticambarus) coosae Hobbs, 1981; C. (P.) extraneus Ha-
gen, 1870; C. (P.) georgiae Hobbs, 1981; C. (P.) hiwasseensis Hobbs, 1981; C.
(P.) parrishi Hobbs, 1981; C. (P.) scotti Hobbs, 1981; Orconectes erichsonianus
(Faxon, 1898); O. forceps (Faxon, 1884); Orconectes spinosus (Bundy, 1877);
Procambarus (Ortmannicus) lophotus Hobbs and Walton, 1960; P. (Pennides)
spiculifer (LeConte, 1856).

Key to Entocytherid Fauna of Northern Georgia
(Based on male copulatory complex)

1. External border of horizontal ramus of clasping apparatus with sim-
DIC-CXCFESCEMCE en4 seagate cen eter up Ankylocythere telmoecea
ly External border of horizontal ramus of clasping apparatus entire .. 2

2(1’). Penis with prostatic and spermatic elements distinctly diverging along
part of their length; ventral part of peniferum deeply cleft posteriorly
Lai ieee Rice =” MUNN MeN Wan toni eon Oot vel het, Meme he Ascetocythere bouchardi
De. Penis with prostatic and spermatic elements contiguous throughout
their lengths; ventral part of peniferum never deeply cleft posteriorly

Saget, MRP os. cucsy $104 tn ta Mahe Noite SaasdecescA ab tig os cheoyl'-\ ott d te eee a oe 3
3(2'). Ventral part of peniferum bulbous .............. Cymocythere cyma
Sie Ventral part of peniferum never bulbous ..................... 2 og
4(3’); «aFinger suard*absent) sg. wih nk oe ee eee 5
4’. Finger guard=presenty.s he Sle ee ee ee Dactylocythere 8
5(4’). Rami of clasping apparatus disposed at angle of no more than 70
ESTES Tee ee en pete cee Donnaldsoncythere donnaldsonensis
5 Rami of clasping apparatus disposed at angle of at least 90 degrees 6
6(5'). Clasping apparatus with internal border of horizontal ramus not ser-
rate, bearing maximum of 3 teeth .......... Uncinocythere simondsi
Oo. Clasping apparatus with internal border of horizontal ramus serrate,
Dearing, SECM g. yc curs: Vans oh eee eee eee Entocythere 7
7(6'). Clasping apparatus with external border of junction of horizontal and
vertical rami produced in angle ............ Entocythere illinoisensis
Teed Clasping apparatus with external border of junction of horizontal and
vertical rami broadly rounded ................. Entocythere elliptica

8(4’). Clasping apparatus “‘U’’ or ‘‘C’’ shaped, distal part expanded
le iS RRA, Be LUNs EAE 0 ee Dactylocythere leptophylax

VOLUME 95, NUMBER 2 305

oe Clasping apparatus ‘*L’’ shaped, distal part tapering ............. 9
9(8’). Accessory groove short, never reaching above dorsal margin of sper-
(TRE JIC ODE Sees aaa 3 Hedi ie wane Lait et cen sp fe <a a 10
oF Accessory groove long, reaching distinctly dorsal to spermatic loop
20 0.5 61" BACB 2 ARIA mele aes Sirona Gel a> Bi lit ty bi. A a a 11

10(9). Accessory groove never reaching dorsal margin of spermatic loop;
vertical ramus of clasping apparatus bowed......................
PN re a. oor. Sala Apel? « Deyo Levey wen ok Dactylocythere brachystrix

10’. Accessory groove reaching dorsal margin of spermatic loop; vertical
ramus sometimes with shoulder but never distinctly bowed
ees. Nd Bec) she Ras bah oe eS Dactylocythere suteri

11(9'). Finger guard jutting anteroventrally ....... Dactylocythere prominula
tele incemeuand directed ventrally] . 2.6.6 28 4 ob. oe aie noe 12

12(11’). External border of junction of horizontal and vertical rami of clasping
apparatus angular; more than one tooth on internal border of hori-
AINA ATINTS) i). nis asain ea Seve sve ee 4 as Dactylocythere mecoscapha

oe External border of junction of horizontal and vertical rami of clasping
apparatus rounded; never more than one tooth on internal border of
MORIZOMAlMATNUS « Adreliioe tnd 2 ee es Dactylocythere falcata

Annotated List of Species

In the following annotated list of ostracods, complete synonymies are furnished
for only those species (Ascetocythere bouchardi and Dactylocythere prominula)
that have been described since the publication of a monograph of the family (Hart
and Hart 1974) in which the remaining species are fully treated. Locality records
are included only for those species that are known from as few as 10 localities.
For information concerning the crayfish hosts, Hobbs (1981) should be consulted.

Ankylocythere telmoecea (Crawford)
Fig. 4

Entocythere telmoecea Crawford, 1959:167-173, figs. 24-30.

Ankylocythere telmoecea.—Hart, 1962:128.—Hart and Hart, 1974:32-33, pl. 4,
figs. 1-5; pl. 46.—Hobbs, 1981:498-501.

Uncinocythere lucifuga.—Hart and Hart, 1974:131 [in part; not Entocythere
lucifuga Walton and Hobbs, 1959].

Range.—‘‘From the Flint River drainage in Georgia to the York Basin in Vir-
ginia’’ (Hobbs and Peters 1977:18).

Previous records in northern Georgia.—Hart and Hart (1974) reported the oc-
currence of Uncinocythere lucifuga in northern Georgia (see locality 4 below).
We have examined the specimen on which this record was based and it appears
to us to be a member of A. telmoecea.

Distribution in northern Georgia.—Except for a single locality in the Chicka-
mauga Creek watershed (Tennessee Basin), all of them are situated in the Etowah
drainage system. Our seven locality records are as follows: CHEROKEE COUNTY:
(1) stream 2 mi W of Free Home on St Rte 20 (hosts: C. (D.) latimanus, P. (Pe.)

306 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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An. telmoecea aa
C. cyma

Dt. brachystrix

Fig. 4. Distribution of Ankylocythere telmoecea, Cymocythere cyma, and Dactylocythere bra-
chystrix in northern Georgia (encircled spot and open triangle denote previously reported localities).

spiculifer; associates: Dt. leptophylax, U. simondsi); (2) stream 1 mi W of Free
Home and 1.5 mi S of Rte 20 (hosts: C.(D.) latimanus, C. (D.) striatus, P. (Pe.)
spiculifer; associate: E. elliptica); (3) Scott Mill Creek at St Rte 140, 2 mi SE of
Canton (hosts: C. (D.) latimanus, P. (Pe.) spiculifer; associates: Dt. falcata, Dt.
leptophylax, E. elliptica, U. simondsi); (4) stream 3.0 mi N of Canton (Hart and
Hart 1974:131) (hosts: C. (D.) latimanus, P. (Pe.) spiculifer; no associate cited).
LUMPKIN County: (5) Etowah River at St Rte 52 (hosts: C. (D.) latimanus, C.
(H.) fasciatus, P. (Pe.) spiculifer; associate: U. simondsi). PAULDING COUNTY:
(6) Possum Creek 1.3 mi NE of U.S. Hwy 278 on St Rte 92 (hosts: C. (D.
latimanus, P. (Pe.) spiculifer; no associates). WALKER COUNTY: (7) trib to Chicka-
mauga Creek 0.5 mi N of St Rte 143 on Rte 341 (hosts: C. (4.) longirostris, C.
(P.) extraneus, O. erichsonianus; associate: Dt. mecoscapha).

Ascetocythere bouchardi Hobbs and Walton
Fig. 5
Ascetocythere bouchardi Hobbs and Walton, 1975:7-10, fig. la—d.

Range.—This ostracod ranges in the Tennessee River basin, on the Cumber-
land Plateau, from Cumberland County, Tennessee southward to Dade County,
Georgia.

VOLUME 95, NUMBER 2 307

Previous records in northern Georgia.—None.

Distribution in northern Georgia.—In Georgia, Ascetocythere bouchardi has
been found in only four localities, all situated on Sand Mountain, Dade County,
in tributaries of the Tennessee River: (1) trib to Warren Creek on unnumbered
road 0.3 mi E of Alabama line and 1.0 mi N of St Rte 301 (host: C. (J.) parvoculus;
no associates); (2) trib to Warren Creek 0.4 mi N of St Rte 301 (hosts: C. (J/.)
parvoculus, C. (J.) unestami; associates: Dt. brachystrix, U. simondsi); (3) creek
at Alabama line about 1.5 mi S of St Rte 301 (hosts: C. VJ.) parvoculus, C. (J.)
unestami; associates: Dt. brachystrix, Dn. donnaldsonensis); (4) Higdon Creek
about 2.0 mi W of Stephensville on St Rte 143 (hosts: C. (J.) parvoculus, C. VJ.)
unestami; associate: Dn. donnaldsonensis).

Cymocythere cyma (Hobbs and Walton)
Fig. 4

Entocythere cyma Hobbs and Walton, 1960:18—19, figs. 11-16.
Cymocythere cyma.—Hart, 1962:129.—Hart and Hart, 1974:44, pl. 9, figs. 7-11;
pl. 47.—Hobbs, 1981:215, 499, 500.

Range.—From Anderson County, Tennessee, to Jefferson County, Alabama,
and from Murray County, Georgia, to Hardin County, Tennessee. Encompassing
the Altamaha, Coosa-Alabama, and Tennessee river basins.

Previous records in northern Georgia.—Only one locality has been cited for
this species in the northern part of the state, this in a small tributary to the
Oostanaula River (Hart and Hart 1974; see below).

Distribution in northern Georgia.—Cymocythere cyma is widely distributed but
uncommon throughout most of the Coosa River basin (including the Chattooga)
of Georgia, although it has not been found in the Etowah watershed. The type-
locality (Four Mile Creek, 1.8 mi SW of Benton, Polk County, Tennessee) is
situated in the Hiwassee drainage system some 20 air miles from the upper reach-
es of the basin in Georgia, but we have no specimens from segments of the river
system in the state. Localities from which our specimens were collected are the
following. FLoyD Country: (1) stream 5.0 mi W of Rome on St Rte 20 (hosts: C.
(D.) striatus, C. (L.) acanthura, C. (P.) coosae, O. spinosus, P. (Pe.) spiculifer;
associates: Dt. falcata, U. simondsi). GILMER COUNTY: (2) Hell’s Creek between
St Rtes 5 and 156, 2.5 mi NW of Pickens Co line (hosts: C. (D.) latimanus, C.
(H.) fasciatus, C. (J.) conasaugaensis, C. (L.) acanthura; associates: Dt. lep-
tophylax, U. simondsi). GORDON COUNTY: 1.8 mi N of Calhoun on U.S. Hwy
41 (Hart and Hart 1974) (hosts: C. (D.) striatus, C. (L.) acanthura, P. (Pe.) spic-
ulifer; no associate cited). MURRAY COUNTY: (4) trib of Conasauga River 0.8 mi
N of St Rte 2 on U.S. Hwy 411 (hosts: C. (D.) striatus, C. (L.) acanthura, C.
(P.) coosae; associates: Dt. falcata, Dt. suteri, U. simondsi); (5) drainage ditch
2.1 mi E of Conasauga River on St Rte 2 (hosts: C. (D.) striatus, C. (L.) acan-
thura; associates: Dt. suteri, U. simondsi). POLK COUNTY: (6) Little Cedar Creek
on Esom Hill Rd, upstream from Hematite Branch (hosts: C. (D.) latimanus, C.
(D.) striatus, C. (P.) coosae; associate: E. elliptica). WALKER COUNTY: (7) 6 mi
S of Lafayette at jct of U.S. Hwy 27 and St Rte 143 (hosts: C. (D.) striatus, C.
(P.) scotti, O. erichsonianus; associate: Dt. falcata).

308 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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Ter ee eel eee 2
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on As. bouchardi a
Dt. falcata

Dt. prominula

Fig. 5. Distribution of Ascetocythere bouchardi, Dactylocythere falcata, and Dactylocythere
prominula in northern Georgia (encircled spot denotes previously reported locality).

Dactylocythere brachystrix Hobbs and Walton
Fig. 4

Dactylocythere brachystrix Hobbs and Walton, 1966:2-3, fig. la-d.—Hart and
Hart, 1974:50-51, pl. 11, figs. 7-11; pl. 48.

Range.—This ostracod is known only from the Tennessee River basin, in which
it ranges on the Cumberland Plateau and Highland Rim from Cumberland County
southwestward to Lawrence County, Tennessee, and on the Plateau and in the
Ridge and Valley Province in Dade and Walker counties, Georgia.

Previous records in northern Georgia.—None.

Distribution in northern Georgia.—Dactylocythere brachystrix has been found
in five localities in the Tennessee Basin on Sand and Lookout mountains. DADE
County: (1) trib to Warren Creek, 0.4 mi N of St Rte 301 (hosts: C. VJ.) par-
voculus, C. (P.) unestami; associates: As. bouchardi, U. simondsi). (2) creek at
Alabama line about 1.5 mi S of St Rte 301 (hosts: C. VJ.) parvoculus, C. (P.)
unestami; associates: As. bouchardi, Dn. donnaldsonensis). (3) trib to Lookout
Creek on St Rte 143, 2.5 mi W of Walker Co line (host: C. (J.) unestami; no
associates); (4) Bear Creek at St Rte 157 (host: C. (J.) unestami; no associates).
WALKER County: (5) Rock Creek at old St Rte 193 on Lookout Mt (host: C.
(J.) unestami; associate: E. elliptica).

VOLUME 95, NUMBER 2 309

Dactylocythere falcata (Hobbs and Walton)
Fig. 5

Entocythere falcata Hobbs and Walton, 1961:379-381, figs. 2, 3, 7, 8.
Dactylocythere falcata.—Hart, 1962:130.—Hart and Hart, 1974:58-S9, pl. 15,
figs. 1-5; pl. 49.—Hobbs, 1981:231, 238, 257, 295, 499.

Range.— ‘From the Coosa River drainage in Georgia and northeastern Ala-
bama to the Tennessee Basin in Virginia, and east of the Appalachian Divide in
the upper James and Roanoke drainage systems in Virginia. The records from
Missouri and Canada cited by Hart and Hart (1974:48, 49) should be confirmed’’
(Hobbs and Peters 1977:30).

Previous records in northern Georgia.—Hart and Hart (1974) reported the oc-
currence of this crayfish in six localities in the Coosa Basin of Georgia: three in
Chattooga (the Floyd County record, 23 mi NW of Rome, is actually in Chattooga
County), and one each in Murray and Whitfield counties (see open circles in Fig.
5) and an unknown one in Walker County.

Distribution in northern Georgia.—This ostracod, one of the commonest in the
area, has been found in 54 localities where it occurs in all of the physiographic
provinces except the Appalachian Plateau and in all of the drainage basins. Hosts
include all of the crayfishes reported from the area except C. (J.) distans, C. (J.)
parvoculus, C. (J.) nodosus, C. (J.) unestami, and O. forceps. It has been found
in association with the following entocytherids: C. cyma, Dt. leptophylax, Dt.
mecoscapha, Dt. suteri, E. elliptica, and U. simondsi.

Dactylocythere leptophylax (Crawford)
Fig. 6

Entocythere leptophylax Crawford, 1961:238-242, figs. 9-14.
Dactylocythere leptophylax.—Hart, 1962:130.—Hart and Hart, 1974:61-62, pl.
16, figs. 1-5; pl. 48.—Hobbs, 1981:70, 257, 450, 598, 500.

Range.—From the Savannah River basin in Oconee County, South Carolina,
westward through the upper Chattahoochee, Ocmulgee, and Coosa basins in
Georgia, and in the French Broad, Little Tennessee, and Hiwassee drainage
systems in Tennessee. A single specimen from the Duck River watershed in
Lewis County, Tennessee (Little Swan Creek in Meriweather Lewis Monument
Park), suggests the occurrence of an outlying population there.

Previous records in northern Georgia.—The only report of the occurrence of
this ostracod in the area is that of Hart and Hart (1974:60) who recorded it from
Neel’s Gap, Union County, on Cambarus (C.) bartonii.

Distribution in northern Georgia.—Dactylocythere leptophylax has been found
in 40 localities in the Ridge and Valley, Blue Ridge, and Piedmont provinces of
northern Georgia where it occurs in the Conasauga, Coosawattee, Hiwassee,
Little Tennessee, and upper Etowah basins. Its hosts in the area are C. (C.)
bartonii, C. (D.) latimanus, C. (H.) coosawattae, C. (H.) fasciatus, C. (H.)
speciosus, C. (J.) conasaugaensis, C. (P.) coosae, C. (P.) hiwasseensis, C. (P.)
parrishi, and P. (Pe.) spiculifer. The entocytherid associates are An. telmoecea,

310 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Distribution of Dactylocythere leptophylax and Dactylocythere mecoscapha in northern
Georgia (encircled spot and open triangle denote previously reported localities).

C. cyma, Dt. falcata, Dt. mecoscapha, Dt. prominula, Dn. donnaldsonensis, E.
elliptica, and U. simondsi.

Dactylocythere mecoscapha (Hobbs and Walton)
Fig. 6

Entocythere mecoscapha Hobbs and Walton, 1960:19-23, figs. 17-20.
Dactylocythere mecoscapha Hart, 1962:130.—Hart and Hart, 1974:62-63, pl. 17,
figs. 1-5; pl. 48.—Hobbs, 1981:380, 499, 500.

Range.—This ostracod occurs in the tributaries of the westwardly flowing seg-
ment of the Tennessee River from Polk County, Tennessee, and Dade, Walker,
Catoosa, and Union counties, Georgia, to Colbert and Lauderdale counties, Al-
abama. The record from Hampton County, South Carolina, cited by Hart and
Hart (1974:63), is almost certainly erroneous, for this low-lying area of the Coastal
Plain Province furnishes unlikely habitats for any member of the genus Dactylo-
cythere except Dt. suteri.

Previous records from northern Georgia.—This ostracod was reported by Hart
and Hart (1974:63) from the Chickamauga National Military Park near Rossville
(Chickamauga Creek basin). Its hosts were P. (O.) lophotus, O. erichsonianus,
and C. (D.) striatus.

Distribution in northern Georgia.—In Georgia, Dactylocythere mecoscapha

VOLUME 95, NUMBER 2 Sil

has been found in 17 localities, all except one of which occurs in the Appalachian
Plateau and Ridge and Valley provinces. There are two records in the Blue Ridge
Province (Wolf Creek and Nottely River) in Union County, so close together as
to be recorded herein by a single spot on Map 6. All 17 localities are in tributaries
of the Tennessee River. In the area under investigation, the hosts of this ostracod
are: C. (C.) bartonii, C. (D.) latimanus, C. (D.) striatus, C. (H.) girardianus, C.
(H.) longirostris, C. (J.) unestami, C. (P.) extraneus, C. (P.) hiwasseensis, O.
erichsonianus, O. spinosus, and P. (O.) lophotus. Its entocytherid associates are
An. telmoecea, Dt. falcata, Dt. leptophylax, Dn. donnaldsonensis, E. elliptica,
E. illinoisensis, and U. simondsi.

Dactylocythere prominula Hobbs and Walton
Fig. 5

Dactylocythere prominula Hobbs and Walton, 1977:606—609, fig. 3.

Range.—Except for the single new locality cited herein, this ostracod is known
to occur only in the Tennessee River basin from Hawkins County, Tennessee,
southward to Walden Gorge and a single outlying locality in the Elk River wa-
tershed in Lincoln County, Tennessee. In this survey, it was found in a tributary
to the Conasauga River in extreme northeastern Murray County, Georgia.

Previous records in northern Georgia.—None.

Distribution in northern Georgia.—Dactylocythere prominula was found in-
festing C. (C.) bartonii in a small unnamed tributary to the Conasauga River
(Blue Ridge Province) about 4.0 mi E of U.S. Hwy 411 on St Rte 2. At this
locality it shared its host with Dt. leptophylax and E. elliptica.

Dactylocythere suteri (Crawford)
Fig. 7

Entocythere suteri Crawford, 1959:162-167, pl. 3.

Dactylocythere suteri.—Hart, 1962:131.—Hart and Hart, 1974:72-73, pl. 21, figs.
11-15; pl. 49.—Peters, 1975:28, figs. 4a, 9—Hobbs and Peters, 1977:41-43, fig.
20, map 10.—Hobbs, 1981:231, 295, 498-500.

Range.—From the Susquehanna Basin in York County, Pennsylvania, south-
ward on the Atlantic slope to the Oconee River drainage in Morgan County,
Georgia. In the Gulf watershed, it occurs in the Conasauga Basin in Murray
County, Georgia, and in the Tennessee Basin in McMinn County, Tennessee.

Previous records in northern Georgia.—Hart and Hart (1974:73) reported the
presence of this ostracod in Holly Creek, Murray County (see below).

Distribution in northern Georgia.—In this area Dactylocythere suteri appears
to be restricted to the Ridge and Valley Province where it has been found in the
following localities in the Conasauga watershed. MurrRAY County: (1) Holly
Creek 1 mi E of Chatsworth on U.S. Hwy 76 (Hart and Hart 1974:73) (hosts:
Cambarus (D.) striatus, C. (P.) coosae, and O. spinosus; associates: Dt. falcata
and U. simondsi); (2) trib to Conasauga River 0.8 mi N of St Rte 2 on U.S. Hwy
411 (hosts: C. (D.) striatus, C. (L.) acanthura, and C. (P.) coosae; associates:
C. cyma, Dt. falcata, and U. simondsi); (3) trib to Conasauga River on St Rte
225 between jct of St Rtes 2E and 2W (hosts: C. (D.) striatus and C. (P.) coosae;

3 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

—F~ Serer ees

=
sl Dt. suteri

SSS
G SaaS

Dn. donnaldsonensis a a Bese
[ee
SSS

E. illinoisensis

Fig. 7. Distribution of Dactylocythere suteri, Donnaldsoncythere donnaldsonensis, and Ento-
cythere illinoisensis in northern Georgia (encircled spot and open triangle denote previously reported
localities).

associate: U. simondsi); (4) drainage ditch 2.1 mi E of Conasauga River on St
Rte 2 (hosts: C. (D.) striatus and C. (L.) acanthura; associates: C. cyma and U.
simondsi); (5) stream 0.2 mi W of St Rte 225 on Rte 286 (hosts: C. (H.) coosae
and O. spinosus; associates: Dt. falcata and E. elliptica).

Donnaldsoncythere donnaldsonensis (Klie)
Fig. 7

Entocythere donnaldsonensis Klie, 1931:334-341, figs. 1-9.

Donnaldsoncythere donnaldsonensis.—Hart, 1962:131-132.—Hart and Hart,
1974:79-80, pl. 23, figs. 11-14; pl. 50.—Hobbs and Peters, 1977:43-45, fig. 21,
map 4 [including a complete synonymy].—Hobbs, 1981:70, 156, 231, 450, 498—
500.

Entocythere hiwasseensis.—Hobbs, 1981:70.

Donnaldsoncythere hiwasseensis.—Hobbs, 1981:70, 156, 231, 450.

Range.—‘‘From northern Georgia to Indiana and Maine’’ (Hobbs and Peters
1977:44).

Previous records in northern Georgia.—Hart and Hart (1974:79) reported it as
Donnaldsoncythere hiwasseensis from Shoal Creek in the Etowah Basin in Daw-
son County where it infested ‘‘C. (Puncticambarus) sp.’ (probably C. (H.) fas-
ciatus).

VOLUME 95, NUMBER 2 318

Fig. 8. Distribution of Entocythere elliptica in northern Georgia (encircled spots denote previously
reported localities).

Distribution in northern Georgia.—Donnaldsoncythere donnaldsonensis has
been found in 22 localities in the Appalachian Plateau, Blue Ridge, and Piedmont
provinces in tributaries of the Tennessee, Hiwassee, and Little Tennessee rivers,
and in headwaters of the Etowah River. Possible hosts in the area are C. (C.)
bartonii, C. (D.) striatus, C. (H.) fasciatus, C. (H.) girardianus, C. (J.) cona-
saugaensis, C. (J.) distans, C. (J.) nodosus, C. (J.) parvoculus, C. (J.) unestami,
C. (P.) georgiae, C. (P.) hiwasseensis, C. (P.) parrishi, and O. erichsonianus.
The entocytherid associates are As. bouchardi, Dt. brachystrix, Dt. leptophylax,
Dt. mecoscapha, E. elliptica, and U. simondsi.

Entocythere elliptica Hoff
Fig. 8

Entocythere elliptica Hoff, 1944:345-349, figs. 15-21.—Hart and Hart, 1974:87-
88, pl. 26, figs. 1, 2; pl. 51.—Hobbs, 1981:71, 231, 295, 312, 399, 403, 431, 450,
498-501.

Range.—Hart and Hart (1974:227) depicted the range of this ostracod as ex-
tending from extreme southeastern Texas to South Carolina and the northern part
of the peninsula of Florida, and northward through most of eastern Mississippi,
Alabama, and Georgia.

Previous records in northern Georgia.—Hart and Hart (1974:88) reported this

314 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

species from two localities in Murray and one each in Fannin and Whitfield
counties.

Distribution in northern Georgia.—As noted in Fig. 8, this ostracod has been
found in 54 localities scattered through all of the physiographic provinces of the
region as well as in all of the watersheds except the Little Tennessee River. Its
23 hosts and 11 entocytherid associates in the area are summarized in Tables 2
and 3.

Remarks.—In treating the entocytherid fauna of North Carolina, Hobbs and
Peters (1977) recognized five species of the genus Entocythere in the state. They
were convinced that Entocythere harrisi Peters (1975:32), E. internotalus Craw-
ford (1959:152), and E. reddelli Hobbs and Walton (1968:243) were all repre-
sented, and although some difficulty was encountered in separating them, they
believed that combinations of characteristics of the male and female genitalia and
those of the pectinate prominence at the base of the apical claws of the antenna
of the female provided a means of recognizing the three.

In studying the Georgia representatives of the genus, we have found those
combinations of characters to be unreliable. Whereas specimens assignable to
each are present in the northern Georgia fauna, there are also individuals that
appear to be typical of Hoff's Entocythere elliptica. Both of us were surprised
when in one specimen of the genus we found one of the paired clasping apparatus
of the male to be typical of that of E. internotalus and the other characteristic of
_E. elliptica. A similar asymmetry of the corresponding elements resembling those
of E. internotalus and E. reddelli was encountered later in several specimens.

Employing the key we had prepared to the North Carolina fauna, we found
that among the specimens from one of the localities in Georgia all of the males
were members of E. internotalus but the females appeared to belong to E. red-
delli. Repeated examination of the material available to us from northern Georgia
has almost convinced us that insofar as the development and position of the
‘‘internal talon’’ on the clasping apparatus is concerned, much of the difference
noted results from the position of the apparatus when viewed, and whereas the
position and development of the teeth (often appearing to be absent) of the talon
may be variable, the direction from which they are viewed minimizes or enhances
the marginal structures of the talon.

The female genitalia are indeed highly variable regardless of the angle at which
they are examined, even more so than those illustrated by Hobbs and Peters
(1977:figs. 25f, 26e, and 27f). The presence of a vermiform tube in E. harrisi, a
simple sinuous sclerotized thickening in E. reddelli, and the compound sclerotized
element in E. internotalus does not characterize the triunguis females collected
in northern Georgia. In at least three of them there are no discernible genitalic
structures. Among other specimens the genitalia are intermediate in their com-
plexity between those of E. internotalus and E. reddelli. Finally, a vermiform
tube was observed in at least one of the females from the Coosa Basin to be
incorporated in a genital complex that is otherwise typical of E. internotalus.

The pectinate prominence at the base of the apical claws of the antenna of the
triunguis female from northern Georgia surprisingly varies from two to at least
10 pectins. Generally the fewer the number the more prominent the pectin. In
seven specimens from a single locality in Cherokee County the full range of
variability was represented, and even in localities in which the number and po-

VOLUME 95, NUMBER 2 315

Table 4.—Variation in size of Entocythere elliptica in the Coosa Basin of northern Georgia.

Length in mm Height in mm

Sub-basin Sex Number Range Average Range Average
Conasauga 3 aD, 0.434-0.490 0.473 0.189-0.210 0.197
2 32 0.560-0.651 0.590 0.273-0.343 0.280

Coosa 3 17 0.413-0.511 0.447 0.175-0.231 0.199
2 16 0.546—-0.630 0.578 0.266—-0.294 0.281

Coosawattee 3 14 0.420-0.483 0.450 0.182—0.224 0.199
2 18 0.532-0.616 0.571 0.245—0.294 0.266

Etowah 3 23 0.406—0.476 0.435 0.175-0.210 0.193
2 24 0.504—0.630 0.548 0.245-0.315 0.278

Entire upper Coosa Basin 3 76 0.406—-0.511 0.450 0.175-0.231 0.197
¢ 90 0.504—0.651 0.577 0.245-0.343 0.276

sition of the pectins exhibit little variability, always one or two individuals
were, in this respect, atypical of the remainder of the population.

In view of the range of the complex, from Texas to Virginia (Hobbs and Peters’
1977:52, 55), and the broad areas within it from which no specimens are available,
we are loath to synonymize any of the three with Hoff’s E. elliptica. Because we
are uncertain as to the identity of the highly variable populations in northern
Georgia, however, we are tentatively assigning our specimens from the area to
the first named of the four, E. elliptica.

Measurements were made of the length and height of the males and females
throughout the study area (Table 4). The average length of specimens from the
Conasauga is greater than that of individuals from the other three sub-basins in
the Coosa watershed and is least in those from the Etowah. In the latter, speci-
mens from the upper part of the drainage system in Dawson and Lumpkin coun-
ties have shorter average lengths (d, 0.429 and 0.417; 2, 0.504 and 0.526 mm,
respectively) than in the middle (d, 0.440; 2, 0.573 mm) and lower parts (d,
0.462; 2, 0.602 mm) of the sub-basin.

Entocythere illinoisensis Hoff
Fig. 7

Entocythere illinoisensis Hoff, 1942:67-69, figs. 1-8.—Hart and Hart, 1974:88—-
90, pl. 26, figs. 3-4; pl. 51.

Range.—From Michigan southward to northwestern Georgia, northern Ala-
bama and Mississippi, and west of the Mississippi River in Arkansas.

Previous records in northern Georgia.—None.

Distribution in northern Georgia.—In this area, Entocythere illinoisensis 1s
known from a single locality, a tributary to Chickamauga Creek 9.8 mi E of the
Dade County line on St Rte 143, Walker County. This stream lies in the Ridge
and Valley Province, and the crayfish hosts were Cambarus (D.) striatus, C. (H.)
girardianus, C. (P.) extraneus, Orconectes erichsonianus, and P. (O.) lophotus.
It was associated with the following entocytherids: Dactylocythere mecoscapha,
Entocythere elliptica, and Uncinocythere simondsi.

316 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

U. simondsi

Fig. 9. Distribution of Uncinocythere simondsi in northern Georgia (encircled spots denote pre-
viously reported localities).

Uncinocythere simondsi (Hobbs and Walton)
Fig. 9

Entocythere simondsi Hobbs and Walton, 1960:17, 18, figs. 1-10.—Hobbs,
1981:70, 71.

Uncinocythere simondsi.—Hart, 1962:138.—Hart and Hart, 1974:133—135, pl. 37,
figs. 4-7, pl. 55.—Hobbs and Peters, 1977:63-64, fig. 33, map 5.—Hobbs,
1981:70, 71, 97, 215, 231, 295, 380, 450, 598-501.

Range.—Hart and Hart (1974:231) reported this crayfish to occur from Illinois
and Kentucky southward to the panhandle of Florida and from Mississippi east-
ward to Georgia and South Carolina. Hobbs and Peters (1977:70) added 25 lo-
calities in the western half of the state of North Carolina.

Previous records in northern Georgia.—This ostracod was reported by Hart
and Hart (1974:134) from seven localities in the area under consideration (Fig. 9,
open circles).

Distribution in northern Georgia.—Uncinocythere simondsi has been collected
in 68 localities in all of the physiographic provinces in the area. Except in the
Nottely and the Hiwassee rivers in Union and Towns counties and in the Little
Tennessee River in Rabun County, it is widespread in northern Georgia where
it has been found on all of the crayfishes in the area except Cambarus (J.) distans,
C. (J.) nodosus, C. (P.) georgiae, C. (P.) parrishi, and Orconectes forceps and

VOLUME 95, NUMBER 2 317

was associated with all of the entocytherids occurring in northern Georgia except
Dactylocythere brachystrix and Dt. prominula.

Acknowledgments

Many persons aided in collecting the specimens on which this report is based.
We are grateful to all of them, and especially to T. A. English, Jr., of Eastern
Air Lines, Atlanta, Georgia, E. T. Hall, Jr., of the Georgia Department of Natural
Resources, and J. E. Pugh, of Christopher Newport College, Virginia, who de-
voted considerable time and effort to assisting us in acquiring the crayfish hosts
from which the ostracods were taken. We extend our thanks to Margaret A.
Daniel for rendering the maps and tables, and to her, T. E. Bowman, and C. W.
Hart, Jr., all of the Smithsonian Institution, for their criticisms of the manuscript.

Literature Cited

Crawford, E. A., Jr. 1959. Five new ostracods of the genus Entocythere (Ostracoda, Cytheridae)
from South Carolina.—University of South Carolina Publications, Biology, series III 2(4):149-
189, 5 plates.

. 1961. Three new species of the genus Entocythere (Ostracoda, Cytheridae) from North and

South Carolina.—American Midland Naturalist 65(1):236-—245, 21 figures.

Hart, C. W., Jr. 1962. A revision of the ostracods of the family Entocytheridae.—Proceedings of the
Academy of Natural Sciences of Philadelphia 114(3):121-147, 18 figures.

Hart, Dabney G., and C. W. Hart, Jr. 1974. The ostracod family Entocytheridae.—Academy of
Natural Sciences of Philadelphia Monograph 18:ix + 239 pages, 49 figures, 52 plates.

Hobbs, Horton H., Jr. 1981. The crayfishes of Georgia.—Smithsonian Contributions to Zoology

318:vili + 549 pages, 262 figures.

, and Daniel J. Peters. 1977. The entocytherid ostracods of North Carolina.—Smithsonian

Contributions to Zoology 247:4 + 73 pages, 33 figures.

, and Margaret Walton. 1960. Three new ostracods of the genus Entocythere from the

Hiwassee drainage system in Georgia and Tennessee.—Journal of the Tennessee Academy of

Science 35(1):17—23, 20 figures.

, and . 1961. Additional new ostracods from the Hiwassee drainage system in Georgia,

North Carolina, and Tennessee.—Transactions of the American Microscopical Society 80(4):

379-384, 8 figures.

, and 1966. A new genus and six new species of entocytherid ostracods (Ostracoda,

Entocytheridae).—Proceedings of the United States National Museum 119(3542): 1-12, 2 figures.

, and 1968. New entocytherid ostracods from the southern United States.—Proceed-

ings of the Academy of Natural Sciences of Philadelphia 120(6):237—252, 3 figures.

, and 1975. New entocytherid ostracods from Tennessee with a key to the species of

the genus Ascetocythere.—Proceedings of the Biological Society of Washington 88(2):5—20,

2 figures.

, and 1977. New entocytherid ostracods of the genus Dactylocythere.—Proceedings

of the Biological Society of Washington 90(3):600-—614, 4 figures.

Hoff, C. Clayton. 1942. The subfamily Entocytherinae, a new subfamily of fresh-water cytherid

ostracods, with descriptions of two new species of the genus Entocythere.—American Midland

Naturalist 27(1):63-—67, 13 figures.

. 1944. New American species of the ostracod genus Entocythere.—American Midland Nat-

uralist 32(2):327-357, 33 figures.

Klie, W. 1931. Campagne spéologique de C. Bolivar et R. Jeannel dans I’ Amérique du Nord (1928),
3: Crustacés Ostracodes.—Biospéologica: Archives de Zoologie Expérimentale et Générale
71(3):333-344, 20 figures.

Peters, Daniel J. 1975. The entocytherid ostracod fauna of the James and York river basins with a
description of a new member of the genus Entocythere.—Virginia Polytechnic Institute and
University, Research Division Bulletin 93:11 + 50 pages, 15 figures.

318 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Walton, Margaret, and Horton H. Hobbs, Jr. 1959. Two new eyeless ostracods of the genus Ento-
cythere from Florida.—Quarterly Journal of Florida Academy of Sciences 22(2):114—120, 20
figures.

(HHH) Department of Invertebrate Zoology, Smithsonian Institution, Wash-
ington, D.C. 20560; (DJP) York High School, P.O. Box 547, Yorktown, Virginia
23690.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 319-324

TWO NEW SPECIES OF LEPTOMYSINID MYSIDS
(CRUSTACEA, MYSIDACEA) FROM
SOUTHERN CALIFORNIA

Linda G. Gleye

Abstract.—Ywo new species of leptomysinid mysids, Cubanomysis myste-
riosa, the second species of the genus described by Bacescu (1968), and Mysi-
dopsis cathengelae are described from the San Onofre region of the Newport-La
Jolla Shelf, California.

Only one species of the genus Cubanomysis is known, C. jimenesi Bacescu,
1968 from Cuba. The species described below has been collected in shallow water,
out to a depth of 8 meters, off the coast of Southern California.

Cubanomysis mysteriosa, new species
Figs. 1-2

Material examined.—Southern California, San Onofre State Beach, 8 meters
depth, 15 January 1977: 3 holotype (USNM 184074), 2 ¢, 2 2 paratypes (USNM
184075).

Etymology.—From the Greek ‘‘mysterion’’ (mystery), referring to the unpre-
dictable occurrence of this species.

Description.—Small, delicate mysid, 3 mm length. Carapace rounded, slightly
produced into rounded rostrum between eyes. Eyes large, globose. Antennal
scale 7 times as long as broad, sparsely setose all around, with large apical seg-
ment, apex bluntly pointed, extending beyond peduncle of antennule by %4 (<)
to '/s (2) its length. Distal joint of antennal peduncle slightly shorter than pre-
ceding joint, whole peduncle extending 7/10 (d) to °/10 (2) length of antennal
scale.

Mandible with large masticatory surface, with small teeth on cutting edge.
Distal segment of palp with 14 plumose setae and 1 terminal simple seta. Maxilla
with small exopod bearing 5 setae.

Endopod of first thoracic limb strong, distal article with dactyl claw and 6
simple setae. Last article of endopod of second thoracic limb with 3 strong setae,
plumose and recurved on the interior face and 7 simple and one plumose setae
on the distal curvature. Endopods of third to eighth thoracic limbs delicate, car-
popropodus divided into 2 subjoints; 8 large articles on exopods.

Sixth abdominal somite twice as long as preceding somite.

First pleopod of adult male without exopod. Pleopods II-V biramous. Pleopod
IV with 2 branches of 6 articles each, exopod longer than endopod, terminating
in a long curved seta bearing 2 rows of fine spines on distal third. Seta extends
past distal tip of telson when pleopod IV is parallel to abdomen.

320 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Cubanomysis mysteriosa: a, Whole body, lateral, 9; b, Rostrum, eyes antennular pedun-
cle, antennal scale, dorsal, ¢; c, Antennal scale and antennal peduncle, ventral 3; d, Right mandibular
palp, 5; e, Maxilla, 6; f, Thoracic endopod 1, distal segments, 3.

Telson short, tip extending to distal end of statocyst, constricted distally. Lat-
eral margins bare. Apex emarginate bearing 16—24 closely set spines.

Endopods of uropods shorter than exopods; inner lower margin with 10 short
spines. Statocysts large with hyaline area appearing swollen.

Remarks.—The present new species is closely allied to C. jimenesi (Bacescu
1968, Brattegard 1969) but differs from the latter in the following features: 1)
more robust form of the endopods of the first and second thoracic appendages,
2) setation of the last article of the endopod of the second thoracic limb, 3)
spination rather than serration of the terminal seta of the exopod of the fourth
pleopod of the adult male and 4) spination of the telson.

There are four known species of the genus Mysidopsis in Southern California,
M. californica Tattersall, 1951, M. onofrensis and M. brattegartii Bacescu and
Gleye, 1969, and M. intii Holmquist (personal observation). The species de-
scribed below has occurred regularly out to a depth of 12 meters in our nearshore
epibenthic samples since the beginning of our studies in 1976.

VOLUME 95, NUMBER 2 S2l

Fig. 2. Cubanomysis mysteriosa: a, Thoracic endopod 2, distal segments, ¢; b, Pleopod 1, 6; c¢,
Pleopod 4, 5; d, Same, terminal seta exopod, distal tip; e, Telson, ¢; f, Uropod, ¢.

Mysidopsis cathengelae, new species
Figs. 3-4

Material examined.—Southern California, San Onofre State Beach, 10 October
1979: 3 holotype (USNM 184076), 2 6,5 2, 1Imm 6 paratype (USNM 18077),
1 ¢,2 2,2Imm ¢ paratype (USNM 184078).

322 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

d

Fig. 3. Mysidopsis cathengelae: a, Whole body, lateral, 2; b, Rostrum, eyes, antennular peduncle,
antennal scale, dorsal, 6; c, Antennular peduncle, antennal scale, antennal peduncle, ventral; 6; d,
Thoracic endopod 1, 2; e, Thoracic endopod 2, 2; f, Thoracic limb 3, 2.

Etymology.—Named for the California zoologist, Dr. Catherine Engel.

Description.—Medium sized, robust mysid, 10 mm. Carapace produced be-
tween eyes into pointed, triangular rostral plate extending nearly to end of eye-
stalks. Eyes of moderate size, cornea occupying half of whole eye in dorsal view.
Antennal scale 9 times as long as broad, narrowly lanceolate, setose all around,
without a distal joint, apex bluntly rounded, extending beyond peduncle of an-
tennule by 7/s (3d) to % (@) of its length. Distal joint of antennal peduncle 7
length of preceding joint, whole peduncle extending less than 12 length of antennal
scale.

VOLUME 95, NUMBER 2 323

DY

Gl
Li] |] J 7

ss a

\

—==S Eg @
[—\ ae
a

——_

Fig. 4. Mysidopsis cathengelae: a, Pleopod 4, 5; b, Telson, 2; c, Uropod, 2. .

Maxillulae with inner lobe small and armed with 2 setae. Maxilla characteristic
of genus, exopod well developed, long and slender, setae present on outer margin,
distal setae much longer than proximal setae.

Endopod of first thoracic limb short and robust, of form characteristic of genus,
dactylus with claw, outer margin armed with 7 heavy and 3 light simple setae.
Endopod of second thoracic limb slightly longer than and less robust than en-
dopod of first limb. Endopod of third thoracic limb longer and more slender than
the second thoracic limb, sixth joint divided into 3 subjoints, second subjoint
shortest, third subjoint longest.

Sixth abdominal somite about 124 times as long as fifth.

Pleopods in male typical for genus, fourth pair with exopod longer than en-
dopod, exopod terminating in long, plumose seta, endopod terminating in two
long, simple setae.

Telson as long as sixth abdominal somite, extending %4 length of inner uropod,
linguiform, apex entire and narrowly rounded. Lateral margins armed with about
40 short spines extending throughout entire length, 5—6 spines distinctly set, re-
maining spines closely set, gradually increasing in length distally, apex armed with
a pair of long spines !/10 length of telson.

Inner margin of inner uropod armed with 12 spines, increasing in length distally,
distal 8 lying more remotely from each other than proximal 4. Exopod of uropod
more than 4 again as long as endopod.

Remarks.—Because of the similarity in telsons, this species may be superfi-
cially confused with M. californica Tattersall. It can be distinguished from the
latter by the following features: 1) larger size, 2) absence of distal suture on
antennal scale, 3) fewer spines on inner margin of the endopod of uropod.

324 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Acknowledgments

I wish to thank Dr. M. Bacescu, Museul de Istorie Naturala “‘Grigore Antipa,”’
for his help with the genus Cubanomysis, and Dr. T. E. Bowman for his help
with M. cathengelae and the preparation of this manuscript.

Funding for this study was supplied by the Marine Review Committee, Inc.

Literature Cited

Bacescu, M. 1968. Etude des quelques Leptomysini (Crustacea Mysidacea) des eaux du Brésil et de

Cuba; Description d’un genre et de cing autres taxons nouveaux.—Annali Museo Civico di

storia Naturale di Genova 77:232-249.

, and L. G. Gleye. 1979. New Mysidacea from the California waters.—Travaux du Museum

d’ Historie Naturelle Grigore Antipa 20:131-141.

Brattegard, T. 1969. Mysidacea from shallow waters in the Bahamas and Southern Florida. Part 1.—
Sarsia 39:17-106.

Tattersall, Walter M. 1951. A Review of Mysidacea of the United States National Museum.—United
States National Museum Bulletin 201:x + 292 pp.

Marine Ecological Consultants, 5333 Stevens Ave., Solana Beach, California
92075. |

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 325-333

THE STATUS OF CIROLANA PARVA HANSEN, 1890
(CRUSTACEA, ISOPODA, CIROLANIDAE) WITH
NOTES ON ITS DISTRIBUTION

Niel L. Bruce and Thomas E. Bowman

Abstract.—The type-material of Cirolana parva Hansen is redescribed, and
the distribution of the species reassessed. Cirolana parva is reliably known only
from the Caribbean Sea and the Gulf of Mexico; most non-Caribbean records are
probably erroneous. Cirolana diminuta Menzies, 1962 is accepted as a valid
species.

Hansen (1890) in his monograph of the Cirolanidae described and figured Cir-
olana parva. All but one of his specimens originated from the West Indies. The
species was later recorded from the Caribbean area (Richardson 1900, 1921;
Moore 1901; Menzies and Glynn 1968) and also throughout the Indo-Pacific and
the East Pacific. The only areas from which the species has never been recorded
are the coasts of South America and Europe.

In examining cirolanids from Australian coasts it became apparent that there
were some inconsistencies amongst specimens being identified as C. parva (Bruce
1980a, 1980b) and Cirolana cranchii australiense Hale 1925 (Holdich et al. 1981).
Detailed study of specimens from selected localities eventually led to the conclu-
sion that what has been considered Cirolana parva is in fact a complex of over
a dozen sibling species, at least 8 of which occur in Australia. It was found to be
necessary to examine the syntypes of Cirolana parva to determine convincingly
that species’ absence from Australia.

Since these species are all so similar to one another we redescribe Cirolana
parva to prevent further application of that name to the numerous closely related,
but distinct species.

Cirolana parva Hansen
Figs, 15 2

Cirolana parva Hansen, 1890:340, pl. 2, figs. 6—6b, pl. 3, figs. 1-1d.—Richardson,
1900:217; 1905:111, figs. 93-95; 1912:178.—Moore, 1901:167, pl. 8, figs. 6-8.—
Menzies & Glynn, 1968:38, fig. 14C—D.

Non Cirolana parva.—Stebbing, 1905:12; 1910:217.—Nobili, 1907:421.—Bar-
nard, 1914:353a; 1936:154; 1940:499.—Chilton, 1924:883, fig. 5; 1926:1980.—
Boone, 1927:136.—Monod, 1930:fig. SA, C; 1931:3; 1933:173, fig. 80; 1937:15.—
Nierstrasz, 1931:151.—Edmondson, 1951:192, fig. 4a~g.—Menzies & Franken-
berg, 1966:51, fig. 27A—C.—Miller, 1968:15, fig 4.—Schultz, 1969:185.—Jones,
1976:216.—Bowman, 1977:653, figs. 1-3.—Hamsa & Nammalwar, 1978:517,
figs. 1-11.—Kensley, 1978:69, fig. 289.—Bruce, 1980a:110; 1980b:158.

Material.—Cirolana parva syntypes: 2 3 (5.6, 6.9 mm), 3 @ (5.1, 5.5, 7.6
mm), St. Thomas, West Indies. 6 (5.9 mm), 2 (6.3 mm) 25°N, 34°E. 2 (7.9 mm,

326 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Cirolana parva syntypes: A—D, F, H, 5.3 mm ¢; E, G, 6.3 mm gd; I-M, 5.5 mm @. A,
Lateral view; B, Cephalon, dorsal view; C, Pleon and pleotelson; D, Pleotelson, posterior border; E,
Peneal processes; F, Clypeal region; G, Sternite 7; H, Pleon, lateral view; I, Pereopod 1; J, Dactyl
of pereopod 1; K, Pereopod 2; L, Antennal peduncle; M, Antennule. Scale line represents 1.0 mm.

ovigerous), West Indies. ¢ (6.9 mm), no locality, West Indies. 2 (5.0 mm) Samoa.
2 specimens (4.4, 5.0 mm), in very poor condition, St. Croix, West Indies. Cir-
olana parva non-types: Specimens from the localities shown in Fig. 4 from the
collections of the National Museum of Natural History, Smithsonian Institution.
Not Cirolana parva: Hundreds of Australian specimens of Cirolana spp., spec-

VOLUME 95, NUMBER 2 327]

Fig. 2. Cirolana parva syntypes: A-D, 5.5 mm 6; E-J, 6.9mm <6. A, Maxilliped; B, Uropod; C,
Uropod exopod, apex; D, Uropod endopod, apex; E, Pereopod 7; F, Appendix masculina, apex; G,
Pleopod 1; H, Pleopod 2; I, Pleopod 4; J, Pleopod 5.

imens of both sexes of C. diminuta Menzies, 1962, from La Jolla, California, and
of C. parva of Jones (1976).

Type-locality.—Hansen (1890) gave no type-locality, but as one of the St.
Thomas specimens has been dissected, it seems apparent that Hansen’s descrip-
tion is based on these specimens. St. Thomas is here recognized as the type-
locality. The type-specimens are held by the Zoologisk Museum, Copenhagen,
Denmark.

Male.—Body about 234 times longer than greatest width, surface minutely
punctate. Cephalon with rostral process; dorsal surface with anterior interocular
carina, and entire faint furrow running between the dorsomedial margin of each
eye, medial part of furrow posteriorly deflected; maxillipedal somite indicated by
2 furrows at posterolateral margins. Pereonite 1 with 2 horizontal furrows; per-
eonites 2-4 each with horizontal furrow, pereonites 5—7 each with oblique furrow;
coxae of pereonites 2—7 each with entire furrow; coxae of pereonites 4—7 pro-
jecting beyond posterior margins of segments. Pleon with segment | largely con-

328 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

cealed by pereonite 7; pleonite 2 with posterolateral margins acute; pleonite 3
with posterolateral margins produced and encompassing pleonite 4, superior mar-
gin convex; pleonite 4 with superior margin rounded, small projection where
horizontal furrow meets margin. Pleotelson with smoothly curving lateral mar-
gins; posterior border with 8 spines, each set within slight identation; short plu-
mose setae lie on each side of spines.

Antennule with peduncular articles 1 and 2 fused; peduncular article 3 with
fused 4th article present; flagellum extending to pereonite 1, composed of 9 ar-
ticles, first of which is longest. Antenna with peduncular articles 1-3 short, pe-
duncular articles 4 and 5 long, subequal in length; flagellum composed of about
26 articles, extending to pereonite 6.

Frontal lamina pentagonal, lateral margins straight, slightly divergent, anterior
margins concave; apex overlapped by downward projection of rostral process.
Clypeus narrow. The fragility of the specimens did not allow a full dissection of
the mouthparts, but they appear similar to those of other members of the genus
(sensu strictu, Bruce 1981). Maxilliped with 2 coupling hooks and 6 plumose setae
on endite.

Pereopod 1 with setae at posterodistal angle of basis; ischium with single spine
at posterodistal angle and 2 setae at superior anterodistal angle; merus with pos-
terior margin slightly sinuate, provided with 5 robust and 2 acute spines; carpus
with single seta and acute spine at posterodistal angle; propodus with 2 spines on
palm, and third stout spine opposing the biunguiculate dactyl. Pereopods 2 and
3 similar, less robust than pereopod | with merus, carpus and propodus propor-
tionally longer, and with more numerous and larger spines. Pereopods 4-7 pro-
gressively longer. Pereopod 7 with virtually no setae, groups of spines at anterior
distal angle of ischium, and propodus with 2 groups of spines, merus and carpus
with one; distal angles of propodus each with 2 spines.

Penes present on sternite 7; short, triangular, projecting posteriorly, set close
together.

Pleopod 1 endopod with slightly concave lateral margin, broadest at 4 of its
length from apex. Pleopod 2 with appendix masculina arising basally, extending
beyond endopod by !/s its length, apex narrows rapidly to an acute slender point.
Exopods of pleopods 3-5 with entire transverse suture. Endopod of pleopod 3
with about 13 setae, pleopod 4 with about 9. Peduncles of pleopods 2-4 each with
4 coupling hooks.

Uropods projecting slightly beyond apex of pleotelson. Exopod slightly shorter
than endopod, lateral margin with 7 spines and short plumose marginal setae,
medial margin with 3 spines; apex bifid with medial process more prominent than
lateral. Endopod with 2 spines and sensory seta on lateral margin; medial margin
with 4 spines amongst plumose marginal setae; apex bifid.

Female.—Differs from the male only in the sexual characters.

Color.—Pale tan in alcohol. Chromatophores bleached out. Related species
show a variety of chromatophore patterns (pers. obs.).

Size.—Syntypes range from 5.6—6.9 mm for the males, and 5.0-7.6 mm for
females.

Variation.—The dorsal cephalic furrow is not always easy to observe and the
indentation of the posterolateral margin of pleonite 4 is only present in the type-

VOLUME 95, NUMBER 2 329

Fig. 3. Cirolana diminuta: A-I, 6.3 mm 6; La Jolla, California. A, Pleon, lateral view; B, Uro-
pods; C, Uropod endopod, apex; D, Uropod exopod, apex; E, Pleotelson, dorsal view; F, Pleotelson,
apex; G, Pleopod 1; H, Pleopod 2; I, Appendix masculina, apex. Cirolana parva, appendix masculina,
apices: J, Montego Bay, Jamaica; K, Quintana Roo, Mexico; L, No Name Key, Florida.

specimens. The apex of the appendix masculina varies slightly as shown in
Fig. 3.

The spination of the pleotelson and uropods, position of the penes, and mor-
phology of pleopods 1 and 2 are constant throughout the specimens examined.

Remarks.—The characters that most readily separate C. parva from other sim-
ilar species are the cephalic furrowing, the spination and shape of the pleotelson
and uropods, the shape of the endopod of pleopod 1, the morphology of the
appendix masculina, the shape and position of the penes, the shape of the lateral
margins of the pleonites and the form of the uropod apices. Cirolana diminuta

330 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Ws 10 105

110 105 100 95 90 85 80 75 70

Fig. 4. Known distribution of Cirolana parva, confirmed from syntypes and from other specimens
in the National Museum of Natural History, Smithsonian Institution. 1, St. Thomas; 2, St. Croix; 3,
San Juan, Puerto Rico; 4, La Parguera, Puerto Rico; 5, Northwestern Cuba; 6, Florida Keys, Florida;
7, Dry Tortugas, Florida; 8, South of Apalachicola, Florida; 9, Montego Bay, Jamaica; 10, Espiritu
Santo Bay, Quintana Roo, Mexico.

can be separated by the differences in the characters figured here (Fig. 3A—L)
and also by having the vasa deferentia opening flush with the surface of sternite
Te

Distribution.—Reliable records exist only for the Caribbean and the Gulf of
Mexico. These are reviewed by Richardson (1905). Menzies and Frankenberg
(1966) record the species from Georgia but figure the uropod with 6 spines on the
medial margin of the endopod, and as the appendix masculina is not quite the
same, this record needs confirmation before it can be accepted. All other records
are either without figures and cannot be confirmed without reference to the spec-
imens, or can be refuted by reference to the figures given (Monod 1930; 1933;
1976), or by reference to specimens (C. parva of Jones 1976, C. diminuta Men-
zies, C. parva of Bruce 1980a, b).

The specimen labelled as from *‘Samoa’’ appears identical to all the other
syntypes, and it is likely that the locality is in error, a possibility considered by
Hansen (1890).

Discussion

In his revision of the family Cirolanidae, Hansen (1890) described Cirolana
parva, the description being accompanied by excellent figures. Several subse-
quent records (Richardson 1900, 1912; Moore 1901) were from the Caribbean, the

VOLUME 95, NUMBER 2 331

source of the original specimens. Shortly after the turn of the century reports
were published from further afield, from Ceylon (Stebbing 1905), the Red Sea
(Stebbing 1910), Polynesia (Nobili 1907) and southeastern Africa (Barnard 1914).
Although none of these early reports gave figures, they apparently led to general
acceptance that Cirolana parva is cosmopolitan in distribution. This acceptance
was reinforced by numerous subsequent records and by 1980 the species had
been recorded from Kenya (Jones 1976), West Africa (Monod 1931), Thailand
(Chilton 1924, 1926), Hawaii (Edmondson 1951; Miller 1968), Indonesia (Nier-
strasz 1931), Cocos Island in the East Pacific (Bowman 1977) and Australia (Bruce
1980a, 1980b). The synonymization of Cirolana diminuta extended the range to
include the western coast of the United States.

In examining “‘Cirolana parva’? from Australia, there appeared to be a great
deal of variation present in samples from one locality and from different localities.
In some series the males had a dense fringe of setae along the inferior margins
of the first pereopod articles. Specimens with this character could not satisfac-
torily be identified as Cirolana parva, yet neither could they be identified as
Cirolana cranchii australiense (see Bruce & Ellis (1982) for discussion of this
species). This prompted the detailed study of all Australian specimens available,
and eventually 8 species were determined. These different species are primarily
separated by differences in details of the pleon, pleopods | and 2, uropod and
pleotelson morphology and spination, setation of the male first pereopod, position
of the vasa deferentia, and articulation and proportional length of the antennule
articles. In general the antenna, mouthparts, pereonal morphology and pereopod
morphology were found ineffective in separating species.

Comparison of the syntypes with specimens recorded by Bruce (1980a) which
include two species, and by Bruce (1980b), and to other unrecorded material
reveals that Cirolana parva does not occur in Australia. Similarly, Kenyan spec-
imens proved not to be C. parva. The record of Hamsa & Nammalwar (1978) is
of a species related to Natatolana woodjonesi (Hale, 1925), and Monod’s (1933)
record, one of the few with figures, is also not C. parva.

One further species needs discussion, Cirolana diminuta, from California.
Menzies and Glynn (1968) concluded that the differences shown by C. diminuta
were insufficient to keep it separate from C. parva. Specimens from La Jolla
identified as C. parva show several differences from C. parva (see Fig. 3), and
the shape of the pleotelson readily identifies them as adult C. diminuta.

A further complication in dealing with the taxonomy of Cirolana parva resulted
from confusion as to whether or not the species was distinct from C. cranchii
Leach. This idea was initiated by Stebbing (1917), and supported by Nordenstam
(1946). Monod (1976) contributed to the argument, and concluded that although
the 2 species were not separable, they were not necessarily conspecific. Bruce
and Ellis (1982) have shown that the 2 species are abundantly distinct, as in fact
can be seen from Hansen’s (1890) figures.

Acknowledgments

The authors thank Dr. J. Just, Universitet Zoologisk Museum, Copenhagen,
for loan of type-material, and Dr. R. C. Brusca, Allan Hancock Foundation, for
donation of Californian specimens. The work was supported by an Australian

387 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Biological Resources Study grant, by an Australian Museum Postgraduate Award,
and by a Commonwealth Postgraduate Study Award.

Literature Cited

Barnard, K. H. 1914. Contributions to the crustacean fauna of South Africa 3. Additions to the
marine Isopoda, with notes on some previously incompletely known species.—Annals of the
South African Museum 10:325a—358a, 359-442.

. 1936. Isopods collected by R.I.M.S. “‘Investigator.’’—Records of the Indian Museum 38: 147—

191.

. 1940. Contributions to the crustacean fauna of South Africa 12. Further additions to the

Tanaidacea, Isopoda and Amphipoda, together with keys for the identification of hitherto

recorded marine and freshwater species.—Annals of the South African Museum 32:381-—S15.

Boone, L. 1927. Scientific results of the first oceanographic expedition of the “‘Pawnee’”’ 1925. Crus-
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Bowman, T. E. 1977. Isopod crustaceans (except Anthuridae) collected on the Presidential Cruise
of 1938.—Proceedings of the Biological Society of Washington 89:653-666.

Bruce, N. L. 1980a. Cirolanidae (Crustacea:Isopoda) of Australia. Heron Island and the Capricorn

Group.—Bulletin of Marine Science 30:108—130.

. 1980b. The Cirolanidae (Crustacea:Isopoda) of Australia: The Coral Sea.—Cahiers de I’ Indo-

Pacifique 2:155-173.

. 1981. The Cirolanidae (Crustacea:Isopoda) of Australia. Diagnoses of Cirolana Leach,

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Freshwater Research 32:945—966.

, and J. Ellis. 1982. Cirolana cranchii Leach, 1818 (Crustacea, Isopoda, Cirolanidae) rede-

scribed.—Bulletin of the British Museum (Natural History), Miscellanea (in press).

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5:875-895.

. 1926. The Tanaidacea and Isopoda of Tale Sap (Siam).—Records of the Indian Museum

28:173-185.

Edmondson, C. H. 1951. Some Central Pacific crustaceans.—Occasional Papers of the Bernice P.
Bishop Museum, Honolulu 20: 183-243.

Hale, H. M. 1925. Review of Australian isopods of the cymothoid group Part 1.—Transactions of
the Royal Society of South Australia 49:128-185.

Hamsa, K. M. S. A., and P. Nannalwar. 1978. Description of isopod Cirolana parva parasitic on the
eyeball of dolphin, Delphinus delphis with a key to the Indian species of the genus Cirolana.—
Journal of the Bombay Natural History Society 75:516—519.

Hansen, H. J. 1890. Cirolanidae et familiae nonnulae propinquae Musaei Hauniensis.—Kongeligt
Videnskabernes Selskab Skrifter, 6, Raekke, Naturvidenskabelig og Mathematisk Afdeling
5:237-426.

Holdich, D. M., K. Harrison, and N. L. Bruce. 1981. Cirolanid isopod crustaceans from the Towns-
ville region of Queensland, Australia, with descriptions of six new species.—Journal of Natural
History 15:555-605.

Jones, D. A. 1976. The systematics and ecology of some isopods of the genus Cirolana (Cirolanidae)
from the Indian Ocean region.—Journal of Zoology, London 178:209-222.

Kensley, B. 1978. Guide to the marine isopods of Southern Africa.—South African Museum, Cape
Town, 173 pp.

Menzies, R. J. 1962. The marine isopod fauna of Bahia de San Quintin, Baja California, Mexico.—

Pacific Naturalist 3:337-348.

, and D. Frankenberg. 1966. Handbook on the common marine isopod crustaceans of Geor-

gia.—University of Georgia Press, Athens, 93 pp.

, and P. W. Glynn. 1968. The common marine isopod Crustacea of Puerto Rico.—Studies on

the Fauna of Curacao and other Caribbean Islands 27:1-133.

Miller, M. A. 1968. Isopoda and Tanaidacea from buoys in coastal waters of the continental United

VOLUME 95, NUMBER 2 353

States, Hawaii, and the Bahamas (Crustacea).—Proceedings of the United States National
Museum 125 (3652):1-53.

Monod, T. 1930. Contribution a |’étude des ‘‘Cirolanidae.’’—Annales des Sciences Naturelles, Zool-

ogie 10(13):129-183.

. 1931. Sur quelques crustacés aquatiques d’ Afrique (Cameroun et Congo).—Revue de Zool-

ogie et Botanie Africaines 21:1-36.

. 1933. Mission Robert-Ph Dollfus en Egypte: Tanaidacea et Isopoda.—Mémoires de 1’ Institute

d’Egypte 21:161—-264.

. 1937. Missions A. Gruvel dans le Canal de Suez. I. Crustacés.—Mémoires de 1’Institute

d’Egypte 34:1-19.

. 1976. Remarques sur quelques Cirolanidae (Crustacea: Isopoda).—Bulletin du Muséum Na-

tional d’ Histoire Naturelle, Paris, 3 série 358, Zoology 251: 133-161.

Moore, H. F. 1901. Report on Porto Rican Isopoda.—Bulletin of the United States Commissioner
of Fish and Fisheries 1900, 20:161-176.

Nierstrasz, H. F. 1931. Die Isopoda der Siboga-Expeditie. 3. Isopoda Genuina 2. Flabellifera.—
Siboga Expeditie, Monographie 32c:123-—233. pls. 10, 11.

Nobili, G. 1907. Ricerche sui crostacei della Polinesia. Decapodi, stomatopodi, anisopodi e isopodi.—
Memorie della Reale Accademia delle Scienze di Torino, serie 2, 57:351-430, pls. 1-3.
Richardson, H. 1900. Synopses of North American invertebrates. VII. The Isopoda.—American

Naturalist 34:207—230, 295-309.

. 1905. A monograph on the isopods of North America.—Bulletin of the United States National

Museum 54:I-LIII, 1-727.

. 1912. Marine and terrestrial isopods from Jamaica.—Proceedings of the United States Na-

tional Museum 42(1894): 178-194.

Schultz, G. A. 1969. How to know the marine isopod crustaceans.—Wm. C. Brown Company Pub-
lishers, Dubuque, Iowa, vii + 359 pp.

Stebbing, T. R. R. 1905. Report on the Isopoda collected by Professor Herdmann at Ceylon, in 1902

in Report to the Government of Ceylon on the Pearl Oyster Fisheries in the Gulf of Manaar,

by W. A. Herdmann, Part 4, Supplementary Report 23:1-64, pls. 1-12.—Royal Society Lon-

don.

. 1910. Report on the marine biology of the Sudanese Red Sea. XIV. On the Crustacea Isopoda

and Tanaidacea.—Journal of the Linnaean Society, Zoology 31:215—230, pls. 21-23.

. 1917. South Africa Crustacea (Part IX. of the S. A. Crustacea, for the Marine Investigations

of South Africa).—Annals of the South African Museum 17:23—-46.

(NLB) Department of Zoology, University of Queensland, St. Lucia, Queens-
land, Australia 4067; (TEB) Department of Invertebrate Zoology (Crustacea),
Smithsonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 334-337

DESCRIPTION OF THE MALE AND NOTES ON
THE FEMALE OF ARGEIOPSIS INHACAE
(CRUSTACEA: ISOPODA: BOPYRIDAE)

Daniel L. Adkison, Richard W. Heard, and Guy T. Clark

Abstract.—The male of Argeiopsis inhacae Kensley, 1974 is described for the
first time from one specimen. The diagnosis for the male is: head fused with first
pereomere; first antenna of 3 segments, second antenna of 5 segments, basal
segment with posteriorly directed projection; midventral tubercles on pereo-
meres; and pleon fused, segments laterally indicated, mid-ventral tubercles, pleo-
pods, and uropods absent. The viability of the subfamily Argeiinae Markham,
1977 is questioned. Based on a review of the literature and examination of the
holotype A. inhacae and specimens from the Philippine Islands, the range of A.
inhacae is extended from Mozambique to the Philippine Islands.

The genus Argeiopsis Kensley, 1974 is based on a single species, Argeiopsis
inhacae Kensley, 1974, described from a single ovigerous female collected from
Mozambique on the banded coral shrimp Stenopus hispidus (Oliver). The present
report is based on the examination of 2 females and a male of A. inhacae obtained
by one of us (GTC) from an aquarium shop in Norfork, Virginia. Host shrimp
were collected in the Philippine Islands and shipped to Norfork, Virginia as part
of a consignment to a marine aquarium shop. Efforts to learn more precise col-
lection data were unsuccessful.

Argeiopsis inhacae Kensley, 1974
Fig. 1

Argeiopsis inhacae Kensley, 1974:259-261, fig. 1.—Markham, 1977:109-—110.

Material examined.—Infesting Stenopus hispidus. Inhaca Island, Mozambique;
from right branchial chamber of Stenopus hispidus; 1 2 (holotype, ovigerous)
SAM A10979.—Philippine Islands; 1977 (aquarium trade, no other collection data
known); from left branchial chamber of S. hispidus; 1 2 (ovigerous), 1 6 USNM
172471. Philippine Islands; 1977 (aquarium trade, no other collection data); from
right branchial chamber of S. hispidus; 1° (non-ovigerous, no ¢) USNM 172472.

Description.—Female. Distortion slight. Head pentagonal, width less than
twice length; anterior lamina present, medially indistinct, laterally curled dorsally.
Small eyes on anterolateral margin of head. First antenna of 3 segments; second
antenna of 4 segments. Maxilliped covered with scales; palp present, medially
with fringe of long setae. Posteroventral lamina with 2 pairs of sickle shaped
projections.

Pereon broadest at pereomere 3; pereomere 3 nearly straight, pereomeres 4 to
7 progressively directed more posteriorly. Narrow coxal plates on pereomeres
1 to 4. Dorsal bosses present on pereomere 1 to 4. Pereomere 1 to 4 bilobed
laterally, tergal area of pereomeres of expanded side enlarged; pereomere | on
enlarged side with tergal area L-shaped or hooked anteriorly. Pereopods increas-

VOLUME 95, NUMBER 2 335

Fig. 1. Argeiopsis inhacae, female a-d (USNM 172472), male e-—h (USNM 172471): a, Dorsal
view; b, Left oostegite 1, internal view; c, Pleon, dorsal view; d, Uropods and right pleopod 5, ventral
view; e, Dorsal view; f, Right antennae; g, Left pereopod 1; h, Pleon, ventral view.

ing in size posteriorly. Oostegites only laterally defining brood pouch. Oostegite
1 with internal ridge unornamented; posterior plate developed into laterally di-
rected point.

Pleon of 6 distinct segments, length subequal to width. Lateral plates absent.
Pleopods biramous, 5 pairs; larger on expanded side; pleopodal endopods de-

336 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

creasing in size posteriorly. Uropods biramous, uropodal exopod similar in size
and shape to exopod of pleopod 5; endopods reduced, smaller than endopod of
pleopod 5.

Variation.—Variation was found in the relative development of coxal plates,
dorsal bosses, pleopodal endopod, and uropodal endopods. The holotype has
weakly developed coxal plates, dorsal bosses and uropodal endopods. On the
other Philippine specimens, these structures are better developed. The uropodal
endopod of the holotype is reduced to a small median projection. The uropodal
endopods of the non-ovigerous female are about of the endopod of pleopod 5S.

Male.—Head narrower than first pereomere; head fused with first pereomere.
Eyes present. First antenna of 3 segments; second antenna of 5 segments, prox-
imal segment with posteriorly directed lobe; second antenna less than twice length
of first; both antennae covered with scales.

Pereon of 7 segments; pereomeres | and 7 narrowed, seventh narrowest; dorsal
pigment spots on pereomere 2 to 7. Pereopods of similar size; covered with scales,
scales with fringe of setae. Midventral tubercles on all pereomeres.

Pleon fused, segments indicated laterally, first most prominent; 5 segments
indicated on right side, 4 on left side. Pleopods, midventral tubercles and uropods
absent. Anus on slight ventral lobe, flanked laterally with setae.

Remarks.—Both the holotype and the ovigerous female from the Philippines
have the cuticle pulled away from the body. This condition makes the determi-
nation of the segmentation of the pleon and pleopods most difficult even when
the specimens are stained.

For female bopyrids with pleopods set on or near the lateral margin, it is
difficult to determine whether one is dealing with a biramous pleopod or a uni-
ramous pleopod and lateral plate. The pleopods of Argeiopsis inhacae are such
a case, and have been considered both uniramous with lateral plate (Markham
1977) and biramous without lateral plate (Kensley 1974). We believe that the
pleopods should be considered biramous (lateral plate absent) since the point of
pleopod attachment is on the ventral surface. We also feel that the presence of
biramous uropods supports this conclusion because the uropods are similar in
structure to the pleopods.

The generic diagnosis presented below is amended from Kensley (1974) to
include the male and changes necessary from the study of the 3 female specimens.

Diagnosis.—Female: Body nearly circular; all regions and segments distinct;
anterior lamina present, not greatly developed; dorsal bosses on pereomeres | to
4; coxal plates narrow or reduced, present on pereomeres 1 to 4; oostegites only
laterally defining brood pouch; pleon without lateral plates; pleopods 5 biramous
pairs; uropod biramous, endopod reduced. Male: head fused with first pereomere;
head narrower than first pereomere; pereon of 7 segments, sides nearly parallel;
pleon with lateral indication of pleomeres; pleon narrower than pereon; pleopods,
midventral tubercles, and uropods absent.

The 5 important differences listed by Markham (1977) distinguishing the Ar-
geiinae from the Bopyrinae (head shape, body outline, pleomere fusion, lateral
plate and uropod development, and number of pleopod rami) appear variable and
insufficient to maintain Argeiinae as a separate subfamily. In addition, the genera
Argeiopsis and Stegoalpheon Chopra, 1923 appear to have biramous pleopods.
Argeiopsis has biramous pleopods and no lateral plates. Chopra (1923), when

VOLUME 95, NUMBER 2 $3)7/

describing Stegoalpheon kempi Chopra, 1923, reported the pleopods to be tri-
ramous (lateral plates absent) but noted that the pleopods would be biramous
were lateral plates present. Of the 5 genera in Argeiinae (Argeia Dana, 1853;
Argeiopsis, Bopyrosa Nierstrasz and Brender a Brandis, 1923; Parargeia Hansen,
1897 and Stegoalpheon), only Parargeia has a closed brood pouch and therefore
appears more similar to the Pseudioninae than to the Bopyrinae. The 4 other
genera have an open brood pouch and are therefore more similar to the Bopyri-
nae. Since the location of the types of Bopyrosa phryxiformis Nierstrasz and
Brender a Brandis, 1923 is unknown and the single known specimen appears to
be immature, its placement is at best tentative (Markham 1977). Until members
of Parargeia and Stegoalpheon can be examined, we tentatively continue to
recognize Argeiinae as a distinct subfamily.

Acknowledgments

We wish to thank Elizabeth Louw of the South African Museum for loan of
the holotype of A. inhacae and Brian Kensley of the National Museum of Natural
History for comments and suggestions offered during a visit to the museum. This

paper is a contribution of the Marine Environmental Sciences Consortium,
P.O. Box 386, Dauphin Island, AL 36528.

Literature Cited

Chopra, B. 1923. Bopyrid isopods parasitic on Indian Decapoda Macrura.—Records of the Indian
Museum 25:411—550.

Dana, J. D. 1853. Crustacea.—United States Exploring Expedition during the years 1838-42 under
the command of C. Wilkes 13(2):691-1018.

Hansen, H. J. 1897. Reports on the dredging operations off the west coast of Central America...
by the U.S. Fish Commission Steamer ‘‘Albatross’’ during 1891 ... XXII. The Isopoda.—
Bulletin of the Museum of Comparative Zoology, Harvard College 31(5):96-129.

Kensley, B. 1974. Bopyrid Isopoda from southern Africa.—Crustaceana 26(3):259-266.

Markham, J. C. 1977. Description of a new Western Atlantic species of Argeia Dana with a proposed
new subfamily for this and related genera (Crustacea Isopoda, Bopyridae).—Zoologische Me-
dedelingen 52(9):107-123.

Nierstrasz, H. F., and G. A. Brender a Brandis. 1923. Die Isopoden der Siboga—Expedition II.
Isopoda Genuina. I. Epicaridea.—Siboga-Expeditie Monographie 32b:57-121, pls. 4-9.

(DLA) Dauphin Island Sea Laboratory, Box 386, Dauphin Island, Alabama
36528 (present address: Department of Biology, Tulane University, New Orleans,
Louisiana 70118); (RWH) Gulf Coast Research Laboratory, Ocean Springs, Mis-
sissippi 39564; (GTC) Department of Biology, Old Dominion University, Norfolk,
Virginia 23508.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 338-346

A DIAGNOSIS OF THE HOBBSI GROUP, WITH
DESCRIPTIONS OF CAECIDOTEA TERESAE, N. SP.,
AND C. MACROPROPODA CHASE AND BLAIR
(CRUSTACEA: ISOPODA: ASELLIDAE)

Julian J. Lewis

Abstract.—The Hobbsi Group has emerged as a major assemblage of both epi-
gean and hypogean species found over much of the eastern United States. With
the new diagnosis of the group, 17 species are now tentatively assigned. Caeci-
dotea teresae, a phreatobite from southern Indiana, is described herein as an
addition to the group, and C. ozarkana is synonymized with C. macropropoda.

During his work on subterranean isopods, Steeves made the first attempt to
divide some of the troglobitic species of the family Asellidae into assemblages
which reflected their evolutionary histories: the Stygia (Steeves 1963), Hobbsi
(Steeves 1964) and Cannulus groups (Steeves 1966). Of these, the Cannulus Group
has received the most attention (Lewis 1981). However, with the addition of new
species and the transfer of others, the Hobbsi Group is emerging as a dominant
element among the North American asellids.

The Hobbsi Group is especially interesting from an evolutionary standpoint,
since unlike the Stygia and Cannulus groups, it contains both epigean and hy-
pogean species. Within the group, morphological intergradations exist between
eyeless, unpigmented troglobites, vestigially eyed and pigmented phreatobites,
to epigean species with functional eyes and pigmentation. The following diagnosis
expands that of Steeves (1964).

The Hobbsi Group

Diagnosis of male. —Eyes and pigmentation present or absent. Palmar margin
of pereopod | propus with strong processes. Pleopod 1 longer than pleopod 2;
lateral margin of exopod concave, with weak nonplumose setae, distal margin
with elongate, plumose setae. Pleopod 2, exopod with numerous elongate, plu-
mose setae along margin of distal segment; endopod, basal apophysis well de-
fined; tip with low conical cannula, extending along the axis of the endopod, often
obscured by other terminal processes.

The following species are tentatively assigned to the Hobbsi group: Troglo-
bites—Caecidotea acuticarpa, C. adenta, C. fustis, C. hobbsi, C. macropropoda,
C. nickajackensis, C. packardi, C. reddelli, C. salemensis; Phreatobites—C. ken-
deighi, C. spatulata, C. teresae, C. tridentata; and Epigean—C. brevicauda, C.
dentadactyla, C. kenki, C. scrupulosa.

Discussion.—The most unusual additions to the Hobbsi Group are undoubtedly
the epigean species. Although Williams (1970) did not assign species to groups,
in his cladogram of epigean asellids C. dentadactyla, C. kenki, C. scrupulosa,
and C. brevicauda formed a lineage related to C. montanus and C. nodula. Of
these latter two species, C. nodula appears to have some affinities with the Hobb-

VOLUME 95, NUMBER 2 339

si Group, especially in the morphology of the first pleopod, but the torsion of the
endopod tip is not apparent in other members of the group. Caecidotea montana’s
relationships with the Cannulus Group have been discussed elsewhere (Lewis
1981).

Unlike other epigean Caecidotea, which occur frequently in warm, lotic hab-
itats, the epigean members of the Hobbsi Group occur mostly in cold water
habitats. This is especially true of C. brevicauda, C. kenki, and C. scrupulosa,
which are spring stream inhabitants and facultative cavernicoles.

Three other species deserve mention, C. parva (included by Steeves 1964 in
the original Hobbsi Group), C. bisetus, and C. pilus. Although the endopod tips
of these species are of the type found in members of the Hobbsi Group, the
similarity ends there. The exopod of the first pleopod is ovate, not laterally con-
cave, and weakly setose; the distal segment of the second pleopod bears only 1-
5 setae; and the palmar margin of the gnathopod propus lacks processes. It is
likely that in the future one or more of these species will be assigned to the genus
Lirceolus.

Caecidotea teresae, new species
Figs. 1—2

Caecidotea sp. #2.—Lewis, 1981:582-583, 585.

Material examined.—INDIANA: Floyd Co., Indiana University Southeast
campus, New Albany, drain tile near parking lot below Natural Sciences Building,
19 Nov 1977, J. Lewis, T. Everitt (Lewis), 28 6, 19 2. Same, 12 Dec 1978, J.
Lewis, 5 36, 6 @. Drain tile flowing into small campus pond, near service area,
22 Mar 1980, T. Lewis, 1 6,2 °@.

A 12 mm male, 19 Nov 1977, is the holotype (USNM 189468), the other spec-
imens are paratypes (USNM 189469, USNM 189470, USNM 189471). All of the
material examined has been deposited in the collections of the National Museum
of Natural History, Smithsonian Institution.

Description.—Eyeless, dorsum with light scattered pigmentation, darkest an-
teriorly. Largest male to about 15.5 mm, female to about 9.5 mm. Body slender,
linear, about 6.8x as long as wide in males, 5.1 in females. Margins of head,
pereonites and telson moderately setose. Head about 1.5x as wide as long, an-
terior margin concave, postmandibular lobes moderately produced. Telson about
1.3x as long as wide, sides subparallel, caudomedial lobe moderately produced.

Antenna | of male reaching midlength of last segment of antenna 2 peduncle,
flagellum with up to about 13 segments, esthete formula 3-0-1-0-1; females with
up to about 9 segments, 3-0-1. Antenna 2 about 0.6 length of body, last segment
of peduncle about 1.3x< as long as preceding segment, flagellum of about 95-100
segments.

Mandibles with 4-cuspate incisors and lacinia; distal segments of palp with
plumose spine rows. Maxilla 1, outer lobe with 13 robust apical spines; inner lobe
with 5 plumose setae apically. Maxilliped with 7 retinacula.

Pereopod | of male, propus 1.5 as long as wide, palm with proximal triangular
process (spine in less mature specimens), larger trapezoidal median process sep-
arated by U-shaped cleft from shorter, rectangular, bicuspid distal process; dactyl
flexor margin with weak spines. Female pereopod 1 propus about 1.5x as long

340 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Caecidotea teresae, A from female paratype, B—H from male paratype: A, Habitus, dorsal;

B, Pleopod 2; C, Same, endopod tip, posterior; D, Same, endopod tip, anterior; E, Uropod, dorsal;
F, Maxilla 1, inner lobe; G, Same, outer lobe; H, Pereopod 1.

as wide, processes absent, large proximal spine present; dacty] flexor margin with
about 6 strong spines. Pereopod 4 more robust and spinose in males than females.

Male pleopod 1 larger than pleopod 2; protopod about 0.7 length of exopod,
with 8 retinacula. Exopod about 0.6 as wide as long, lateral margin concave,
with about 11 elongate plumose setae along distal margin. Pleopod 2, protopod

VOLUME 95, NUMBER 2 34]

Fig. 2. Caecidotea teresae, male paratype: A, Head and antennae, dorsal; B, Antenna 1, distal
segments; C, Pereopod 4; D, Pleopod 3; E, Pleopod 1; F, Pleopod 4; G, Pleopod S.

with about 10 setae along medial margin. Exopod, proximal segment with up to
8 lateral setae, distal segment with about 23 elongate, plumose setae. Endopod
with distinct basal apophysis; tip of endopod terminating in 3 processes: (1) can-
nula trapezoidal, low, wide, apically blunt, partly obscured by other processes;
(2) mesial process slightly longer than cannula, slightly recurved, forming lateral

342 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

wall of endopodial groove; and (3) caudal process low, broadly rounded, heavily
sclerotized. Female pleopod 2 pyriform, with about 13 elongate plumose setae
along lateral margin, 2—3 setae on proximal-mesial margin. Pleopod 3 exopod,
proximal segment about 0.7 length of distal segment, with about 12—13 setae on
lateral margin; distal segment with about 19-20 elongate plumose setae on distal
margin. Pleopod 4, about 5 setae on proximal lateral margin, 2 false sutures
present. Pleopod 5, 3 setae on proximal lateral margin.

Uropods showing pronounced sexual dimorphism; male uropod about 1.5x
length of pleotelson; female uropod about 0.7 length of pleotelson.

Etymology.—This species is named for Teresa M. Lewis, in recognition of her
collection of not only this species, but many other subterranean asellids from the
eastern U.S. |

Relationships.—Caecidotea teresae has its closest morphological affinities with
C. tridentata, C. salemensis, and an undescribed species from Illinois (Lewis and
Bowman 1981). The relationship of Caecidotea teresae to these species was dis-
cussed in detail previously (Lewis 1981) and will not be repeated here.

Habitat and distribution.—Caecidotea teresae is known only from drain tiles
on the New Albany campus of Indiana University Southeast. A small stream
flows from a drain tile near the Natural Sciences Building, which at times is
inhabited by a dense population of C. teresae. In addition to the phreatobitic
isopods, an undescribed amphipod of the genus Stygobromus has also been taken
from a drain tile near the campus service center. Both of these drain tile streams
disappear during dry weather.

Caecidotea macropropoda Chase and Blair
Fig. 3

Caecidotea macropropoda Chase and Blair, 1937:220—224.—Nicholas, 1960:131.
Kenk, 1973:13.

Caecidotea ozarkana Chase and Blair, 1937:220—224. Nicholas, 1960:131.

Caecidotea macropropodus.—Mackin, 1940:17-18.—Mackin and Hubricht,
1940:395.

Asellus macropropodus.—Hubricht, 1950:17.—Dearolf, 1953:227.—Bresson,
1955:51.—Mackin, 1959:875.—Black, 1971:6.—Fleming, 1973:294, 298.

Asellus ozarkanus.—Bresson, 1955:51.—Fleming, 1973:294, 298.

Conasellus macropropodus.—Henry and Magniez, 1970:356.

Conasellus ozarkanus.—Henry and Magniez, 1970:356.

Asellus ozarkana.—Black, 1971:7.

Material examined.—OKLAHOMA: Adair Co., stream in cave, 5 mi S. Kan-
sas, A. P. Blair, 27 Dec 1935, 8 6 (USNM 8625); cave (same as previous collec-
tion?), A. P. Blair, 27 Dec 1935, 3 ¢; Spring, 5 mi S. Kansas, A. P. Blair, 12 Jul
1936, 1 6, 3 2 (USNM 108660); Cave, 5 mi. S. Kansas, coll. unknown, 15 Sep
1960, 3 6, 3 2; Bat Cave, 5 mi. S. Kansas, J. Lewis, T. Lewis, 1 June 1981, 17
6,9 2; Cave behind old greenhouse, J. Lewis, T. Lewis, 1 June 1981, 2 6,5
2; Cave behind Hardwicks house, 1 June 1981, 1 6.

Description of male.—Eyeless, unpigmented, length to about 15.0 mm. Body
slender, linear, about 6x as long as wide; coxae visible in dorsal view. Head
about 1.8x as long as wide, anterior margin concave, rostrum absent. Telson
about 1.4x as long as wide, sides subparallel.

VOLUME 95, NUMBER 2 343

Fig. 3. Caecidotea macropropoda, A-—B, from male topotype of C. ozarkana, spring 5 miles south
Kansas; C—H from male topotypes of C. macropropoda, Bat Cave, 5 miles south Kansas: A, Pleopod
2 endopod tip, anterior; B, Same, posterior; C, Same, posterior; D, Pereopod 1, palmar margin of
propodus; E, Antenna 1, distal segments, with abnormal esthete placement on Sth segment; F, Pleo-
pod 1; G, Pleopod 2; H, Pleopod 4. Caecidotea stiladactyla, male, from small seep 9 miles southwest
of Harrison, Boone Co., Arkansas (USNM 108666): I, Pleopod 2, endopod tip, anterior, recumbent
position.

Antenna | of about 13 segments, esthete formula 3-0-1. Antenna 2, last segment
of peduncle about 1.5x length preceding segment, flagellum to about 105 seg-
ments.

Mandibles with 4-cuspate incisors and lacinia mobilis; distal segments of palp

344 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

with plumose spine rows. Maxilla 1, outer lobe with 13 robust spines, inner lobe
with 5 robust, plumose setae. Maxilliped with 4—S retinacula.

Pereopod | propus about 1.5 x as long as wide, with 3 processes: stout proximal
spine, or spine-like process; median process large, blade shaped; distal process
shorter, bicuspid, becoming lower and broader in kouget individuals. Dactyl flexor
margin with massive, lobed process.

Pleopod 1 longer than pleopod 2, protopod with 6-7 retinacula; exopod about
1.8x as long as wide, widened proximally with 1-2 proximomedial setae, lateral
margin concave, bearing tiny spines, distal margin with about 10 non-plumose
setae. Pleopod 2, exopod proximal segment with 4—5 small, lateral setae; distal
segment with about 22 elongate, plumose marginal setae. Endopod with large,
slightly decurved basal apophysis, tip with 3 processes: cannula decurved, tu-
bular, flanked by broadly rounded, heavily sclerotized caudal process, and sim-
ilar, but more slender, subterminal lateral process. Pleopod 3 with about 8 plu-
mose setae along distal margin. Pleopod 4 with 2 false sutures, proximal lateral
setae present. Pleopod 5 with poorly defined transverse suture. Uropods spatu-
late, over 2x length of pleotelson.

Remarks.—The report of 2 troglobitic asellids, Caecidotea ozarkana and C.
macropropoda, from adjacent localities 5 miles south of Kansas, Oklahoma by
Chase and Blair (1937) presented an unusual zoogeographic situation. Although
not unprecedented, the syntopy (or near syntopy, as is the case here) of 2 trog-
lobitic Caecidotea has been reported in only a few instances (Culver 1976; Lewis
and Lewis 1981). Examination of specimens collected by A. P. Blair from the
type localities of the 2 species confirmed the suspicion that C. ozarkana and C.
macropropoda are conspecific. As C. ozarkana was apparently described from
immature specimens taken from an unspecified spring, C. macropropoda 1s pro-
posed here as the senior synonym.

Relationships.—The general morphological affinities of Caecidotea macropro-
poda appear to lie with the members of the Hobbsi Group. The strongly armed
gnathopod and the shape and setation of the first pleopod are typical of other
members of the group, but the structure of the second pleopod endopod tip is
somewhat unusual. The cannula is recurved and appears to twist somewhat on
its axis, in contrast to the usual low, conical cannula found in the Hobbsi Group.
However, the basic shape of the cannula is conical and is obscured by other tip
processes, as would be expected. The placement of Caecidotea macropropoda
in the Hobbsi Group should be considered tentative.

Caecidotea macropropoda is most similar morphologically to C. stiladactyla,
which occurs in caves and springs in the part of Arkansas adjacent to the localities
of C. macropropoda. In both species the cannula appears movable, but in Cae-
cidotea stiladactyla the cannula apparently extends and retracts from the body
of the endopod. For comparison with C. macropropoda, the endopod tip of C.
stiladactyla is illustrated in the retracted position (Fig. 3). Caecidotea macropro-
poda and C. stiladactyla may be easily separated by the morphology of the fourth
pleopod. In C. macropropoda, 2 false sutures are present, ae in C. stiladactyla
only a single, sigmoid suture is present.

Distribution and habitat.—Dearolf (1953) reported Asellus macropropodus
from 3 additional Oklahoma caves, besides the type-localities of C. ozarkana and
C. macropropoda, plus a cave and an adjacent pool in Arkansas. These localities

VOLUME 95, NUMBER 2 345

remain unconfirmed. Besides the cave region of northeastern Oklahoma, which
consists of an extension of the Springfield Plain section of the Ozark Plateau into
the corner of the state, collections from the adjacent parts of Arkansas, Missouri,
and Kansas have also been examined in search of new localities for C. macro-
propoda. However, Caecidotea macropropoda still remains known only from the
type-locality and the immediate vicinity.

The type-locality of Caecidotea macropropoda, Bat Cave, is now also known
as Christian School Study Cave. This cave was visited in June 1981 and found
to be inhabited by a large population of isopods in a guano bog a short distance
inside the lower entrance. The cave is also inhabited by a colony of Gray Bats,
Myotis griscesens, which contribute the guano enrichment to the stream. The
type-locality of C. ozarkana was not specified by Chase and Blair (1937), but
Several small springs and spring caves exist in the same valley as Bat Cave.
Caecidotea macropropoda was also found at 2 of these small caves, along with
numerous planarians, Dendrocoelopsis americana (Kenk 1973). Presumably, one
of these small springs was the type-locality of C. ozarkana.

Black (1971) presented some water quality data for Bat Cave. In comparison
with other Oklahoma caves surveyed, Bat (Christian School Study) Cave had
particularly high total dissolved solid (209 ppm), nitrate (17.0 ppm) and ortho-
phosphate (4.2 ppm) levels, probably reflecting the guano enrichment of the
stream.

Acknowledgments

I would like to thank Murray and Mary Looney for providing information on
Oklahoma caves, field support, and other assistance during a collecting trip
through the Ozarks. Thomas E. Bowman read the manuscript and provided the
loan of specimens from the collections of the Smithsonian Institution. Travel
funds were provided by a grant from the Graduate School of the University of
Louisville. Finally, I thank Teresa M. Lewis for her assistance in making nu-
merous collections in the Ozarks and Ouachitas.

This paper is contribution number 202 (New Series) from the Department of
Biology, University of Louisville.

Literature Cited

Black, J. H. 1971. The cave life of Oklahoma.—Oklahoma Underground 4(1 & 2):2—56.

Bresson, J. 1955. Aselles de sources et de grottes d’Eurasie et d’ Amerique du Nord.—Archives de
Zoologie et Generale 92(2):45-77.

Chase, H. D., and A. P. Blair. 1937. Two new blind isopods from northeastern Oklahoma.—American
Midland Naturalist 18:220-224.

Culver, D. C. 1976. The evolution of aquatic cave communities.—American Naturalist 110:945—957.

Dearolf, K. 1953. The invertebrates of 75 caves in the United States.—Pennsylvania Academy of
Science 27:225-241.

Fleming, L. E. 1973. The evolution of the eastern North American isopods of the genus Asellus
(Crustacea: Asellidae).—International Journal of Speleology 5:283-310.

Henry, J.-P., and G. Magniez. 1970. Contribution a la systematique des Asellides (Crustacea: Isop-
oda).—Annales de Spéléologie 25(2):335—367.

Hubricht, L. 1950. The invertebrate fauna of Ozark caves.—Bulletin of the National Speleological
Society 12:16-17.

Kenk, R. 1973. Freshwater triclads (Turbellaria) of North America, VI: The genus Dendrocoelop-
sis.—Smithsonian Contributions to Zoology 135:1-16.

346 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Lewis, J. J. 1980. A comparison of Pseudobaicalasellus and Caecidotea, with a description of Cae-

cidotea bowmani, n. sp. (Crustacea: Isopoda: Asellidae).—Proceedings of the Biological So-

ciety of Washington 93(2):314—326.

. 1981. Caecidotea salemensis and C. fustis, new subterranean asellids from the Salem Plateau

(Crustacea: Isopoda: Asellidae).—Proceedings of the Biological Society of Washington

94(2):579-S90.

, and T. E. Bowman. 1981. The subterranean Caecidotea of Illinois (Crustacea: Isopoda:

Asellidae).—Smithsonian Contributions to Zoology 335:1-66.

, and T. M. Lewis. 1980. The distribution and ecology of two species of subterranean Cae-

cidotea in Mammoth Cave National Park.—Cave Research Foundation Annual Report

1980:23-27.

Mackin, J. G. 1940. A key to the Oklahoma species of the family Asellidae.—Proceedings of the

Oklahoma Academy of Science 20:17-18.

. 1959. Malacostraca (Isopoda). Pp. 869-901 in Edmondson, W. T., ed., Freshwater Biology,

2nd Ed.—John Wiley and Sons, Inc., New York.

, and L. Hubricht. 1940. Descriptions of seven new species of Caecidotea (Isopoda: Asellidae)

from the central United States.—Transactions of the American Microscopical Society 59:383-

397

Nicholas, G. 1950. Checklist of macroscopic troglobitic organisms of the United States.—American
Midland Naturalist 64(1):123-160.

Steeves, H. R. III. 1963. The troglobitic asellids of the United States: the Stygius Group.—American

Midland Naturalist 69(2):470-481.

. 1964. The troglobitic asellids of the United States: the Hobbsi Group.—American Midland

Naturalist 71(2):445—451.

. 1966. Evolutionary aspects of the troglobitic asellids of the United States: the Hobbsi, Stygius

and Cannulus Groups.—American Midland Naturalist 75(2):392—403.

Williams, W. D. 1970. A revision of North American epigean species of Asellus (Crustacea: Isopo-
da).—Smithsonian Contributions to Zoology 49: 1-80.

Department of Biology, University of Louisville, Louisville, Kentucky 40292.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 347-351

GONODACTYLUS INSULARIS, A NEW STOMATOPOD
CRUSTACEAN FROM ENEWETAK ATOLL,
PACIFIC OCEAN

Raymond B. Manning and Marjorie L. Reaka

Abstract.—The ninth species of the G. falcatus complex in the Indo-West
Pacific region is recognized. Like other species of the complex, it can be distin-
guished in the field by its color pattern.

Until relatively recently, Gonodactylus falcatus (Forskal, 1775) was considered
to be one of the two most widely distributed members of the genus in shallow
water habitats in the Indo-West Pacific region, ranging from the Red Sea and
East Africa eastward to Hawaii and Japan. Field studies conducted by R. L.
Caldwell, University of California, H. Dingle, University of Iowa, and their col-
leagues, including one of us (MLR), strongly suggested that G. falcatus actually
was a complex of species sharing many morphological features as well as habitat
preferences, but differing in size, behavior, and color in life. Reexamination of
_ type-specimens of taxa considered to be junior synonyms of G. falcatus resulted
in the recognition of five species (Manning 1978), and two species have been
described since then (Manning and Reaka 1981a, b). The other member of the
complex, G. graphurus Miers, 1875, from Australasian localities, is distinctive
morphologically (Manning 1978:fig. 2) and has not been confused with other mem-
bers of the complex in the recent literature.

We name here the ninth species of the complex, based upon specimens col-
lected at Enewetak Atoll. All specimens have been deposited in the collections
of the National Museum of Natural History, Washington, under USNM catalogue
numbers.

We thank C. W. Hart, Jr., Smithsonian Institution, and Austin B. Williams,
Systematics Laboratory, National Marine Fisheries Service, for their comments
on the manuscript. Philip Helfrich facilitated field work by one of us (MLR) at
the Enewetak Marine Biological Laboratory in 1972 with a grant from that insti-
tution.

Gonodactylus insularis, new species
Fig. 1

Gonodactylus falcatus.—Manning, 1971:73, 74.—Reaka, 1975:56; 1976:484
[discussion]; 1979:238, 248, 249, 252, figs. 2-4.—Dingle et al., 1973:55, 58, 59,
o2 table 2.

Material.—Kidrenen [Keith or Grinem] Island, Enewetak, 11°22’S0’N,
162°10’30’E, lagoon side, scattered small coral heads on bottom of coral sand at
75 feet, C. A. Child, sta. 7-69, 23 September 1969: 1 female (holotype, USNM
135632).—Enewetak, 1971(?), coral rubble, 0-1 meters, R. L. Caldwell: 2 females
(paratypes, USNM 139883).—Enewetak, Aomon (Sally) Island, coral rubble, 1-

348 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Gonodactylus insularis, female paratype, total length 31 mm: a, Rostral plate and ocular
scales; b, Sixth abdominal somite and telson; c, Uropod, ventral view.

3 meters, 1972, M. L. Reaka: 3 males, 10 females (paratypes, USNM 189383).—
Sand Island, Enewetak, R. L. Caldwell et al., August 1971: 6 males, 5 females
(USNM 189384).

Description.—Rostral plate with length and width subequal or slightly longer
than broad, median spine relatively short, anterior margins of basal part of plate
at right angles to body line or sloping anteriorly, lateral margins divergent, an-
terolateral angles acute but rounded. Ocular scales small, breadth of each no
greater that that of rostral spine at base. Anterior 5 abdominal somites lacking
transverse grooves, sixth somite with 6 carinae, variously inflated, each usually
with strong posterior spine; median carinule absent. Small black spot usually
present on each side of sixth somite between submedian and intermediate carinae.
Abdominal width/carapace length index ranging from 882 in smaller specimens
to 740 in larger ones. Telson appearing elongate because of slender marginal
teeth, length and width subequal or width slightly greater. Dorsal carinae of telson
slender or moderately inflated, median and accessory medians each usually with
strong apical spine flanked ventrally by strong excavation. Knob distinctly bi-
lobed. Three pairs of marginal teeth present, submedians with movable apices;
apices of intermediate teeth slender, sharp, curved mesially; lateral teeth shorter,
blunter. Anterior surface of telson with dark spot on each side anterior to each
anterior submedian carina. Ventral surface of telson with sharp carina on each
submedian tooth. Uropodal exopod with single line of marginal setae, exopod
with 9-12, usually 11, graded movable spines on outer margin of proximal seg-
ment; basal prolongation with one rounded lobe proximally on inner margin of
outer spine.

Color.—Body of both sexes similar, greenish or brownish, heavily speckled or
mottled, lacking red posterior bands on abdominal somites in both sexes. Anten-
nal scale variable in color, clear, red, yellow, orange, yellow-green, or blue-green,
often (but not always) with white, blue-white, or blue patches or speckles, usually
with pink or red edges; two specimens (both females) with base of scale red, and
tip blue or blue-green. Meral spot yellow with anterior reddish infusion, not

VOLUME 95, NUMBER 2 349

Fig. 2. ?Gonodactylus insularis, female, total length 39 mm, Onatoa: a, Rostral plate and ocular
scales; b, Sixth abdominal somite and telson; c, Uropod, ventral view.

variable; smaller flanking spots as follows: dorsal posterior spot blue-green and
black (not variable); dorsal anterior crescent variable, blue-green, brown, or
_ Clear; anterior spot variable, black, red, or brown; ventral spot green or brown,
variable. Other maxillipeds variable in color, green, or blue-green, or usually
yellow or yellow-green with pink front edges. Pereopods variable, sexes essen-
tially similar, merus variably greenish or yellow or pink with white splotches,
distal two segments red or pink; some individuals (both male and female) with
bluish coloration on pereopods. Uropods similar in both sexes, not variable,
endopod and exopod red or pink with white mottling or spots, setae red.

Measurements in mm.—Total lengths of males 13 to 30, of females 10 to 31.5.
Other measurements of female holotype: carapace length 7.3; rostral plate length
2.4, width 2.4; fifth abdominal somite width 5.5; telson length 4.5, width 4.5.

Reaka (1979:252, table 7) noted that at Enewetak this species settled at a length
of 8 mm, was 13 mm long at sexual differentiation, reached reproductive maturity
at 31 mm, and had a maximum size of 34 mm.

Remarks.—This new species resembles three of the species of the G. falcatus
complex that have a bilobed knob on the telson and that lack a median carinule
on the sixth abdominal somite and for which color in life is known: G. aloha
Manning and Reaka, 1981, from Hawaii, and G. mutatus Lanchester, 1903, and
G. siamensis Manning and Reaka, 1981, both from Thailand. The most striking
differences among these species are their color patterns in life. The diagnostic
meral spot in G. insularis is basically yellow, as in these three other species, but
the anterior infusion is reddish whereas it is brown in G. aloha, G. mutatus, and
G. siamensis. Gonodactylus insularis lacks posterior red bands on any of the
abdominal somites in either sex; these are also absent in G. mutatus, but are
present in males of G. aloha and G. siamensis. The uropodal endopod in G.
insularis is red or pink with white spots and red setae; in G. aloha they are
pinkish-orange with yellow distally and green or pink setae. The uropod color has
not been reported in G. mutatus.

350 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Gonodactylus insularis differs from both G. mutatus and G. siamensis in having
narrower Carinae with relatively long posterior spines on the median and acces-
sory carinae on the telson, and much sharper, more slender intermediate marginal
teeth on the telson. These features of the telson are so distinctive that we tentatively
identify with this species two females, 27.5 and 39 mm long, from Onotoa, Gilbert
Islands (Fig. 2). Gonodactylus insularis may well prove to frequent other oceanic
islands in the Pacific.

Nothing is known of the color pattern of G. glabrous Brooks, 1886, from the
Philippines and Indonesia, but it is a larger species, 50 to 51 mm long, and the
teeth and spines of the telson are all slender and sharp (Manning 1978:fig. 3).

The abdominal width/carapace length indices of G. insularis, like those of G.
siamensis, are Somewhat variable, as follows:

Enewetak
(all but Sand Island) Sand Island
Index Index
Carapace No. of ————$
length, mm specimens Range Mean Range Mean
3 3 794-824 814 882 882
4 2 800-806 803 821-875 848
5) 2) 800-820 810 784-849 817
6 3 764-797 783 —= =
7 1 WS) 753 821 821

It seems likely from the abdominal width/carapace length indices given above
that the specimens from Sand Island represent a separate species. Samples from
Sand Island, an apparently temporary emergent island between Enewetak (Fred)
and Parry islands, on the southeastern quadrant of the atoll, were collected by
R. L. Caldwell and colleagues in 1971. When researchers returned to Enewetak
in 1972, Sand Island and its population of Gonodactylus no longer existed.

In the 1971-1972 Annual Report of the Eniwetok Marine Biological Laboratory,
R. L. Caldwell and H. Dingle reported that they found three species of gonodac-
tyloids on Sand Island, one of which was referred to as Gonodactylus sp. Dingle
et al. (1973:58) reported on their collections from 1971, and noted: °“‘G. falcatus,
on the other hand, when we did find it, occupied rubble that was conspicuous for
the attached overgrowing clumps of green algae.’’ They also noted that coral
rubble fragments with attached green algae were uncommon on Sand Island,
possibly a reflection of the scarcity of the species they identified with G. falcatus.

Without information on color in life of the population from Sand Island, we are
reluctant to characterize it as a species distinct from G. insularis proper. In view
of this, we tentatively identify the sample from Sand Island with G. insularis, but
do not consider those specimens as paratypes.

The holotype, the largest specimen available, appears to be conspecific with
the remainder of the type-series. However, unlike the remainder of the speci-
mens, which were collected in relatively shallow water, the holotype was col-
lected at a depth of 75 feet. Whether the species actually occurs from the shallow
intertidal to that depth or whether the holotype washed to that depth is unknown.

Etymology.—The specific epithet is from the Latin, ‘‘insularis’’, of an island.

Literature Cited

Dingle, Hugh, R. C. Highsmith, K. E. Evans, and Roy L. Caldwell. 1973. Interspecific aggressive
behavior in tropical reef stomatopods and its possible ecological significance.—Oecologia
13:55-64.

VOLUME 95, NUMBER 2 pil

Manning, Raymond B. 1971. Two new species of Gonodactylus (Crustacea, Stomatopoda), from
Eniwetok Atoll, Pacific Ocean.—Proceedings of the Biological Society of Washington 84:73-
80, figures 1, 2.

. 1978. Notes on some species of the Falcatus Group of Gonodactylus (Crustacea: Stomato-
poda: Gonodactylidae).—Smithsonian Contributions to Zoology 258:1-15, figures 1-13.

, and Marjorie L. Reaka. 198la. Gonodactylus aloha, a new stomatopod crustacean from the
Hawaiian Islands.—Journal of Crustacean Biology 1(2):190—200, figures 1-3.

, and . 1981b. Gonodactylus siamensis, a new stomatopod crustacean from Thailand.—
Proceedings of the Biological Society of Washington 94(2):479-482, figure 1.

Reaka, Marjorie L. 1975. Molting in stomatopod crustaceans, I. Stages of the molt cycle, setagenesis,
and morphology.—Journal of Morphology 146(1):55—80, figures 1-25.

. 1976. Lunar and tidal periodicity of molting reproduction in stomatopod Crustacea: a selfish
herd hypothesis.—Biological Bulletin 150:468—490, figure 1.

. 1979. The evolutionary ecology of life history patterns in stomatopod Crustacea. Pp. 235-—
260 in S. E. Stancyk (editor), Reproductive ecology of marine invertebrates.—University of
South Carolina Press: Columbia.

(RBM) Department of Invertebrate Zoology, National Museum of Natural His-
tory, Smithsonian Institution, Washington, D.C. 20560; (MLR) Department of
Zoology, University of Maryland, College Park, Maryland 20742.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 352-353

RISSOIDES, A NEW GENUS OF STOMATOPOD
CRUSTACEAN FROM THE EAST ATLANTIC
AND SOUTH AFRICA

Raymond B. Manning and Ch. Lewinsohn

Abstract.—A new genus is recognized for 5 East Atlantic species formerly
assigned to Meiosquilla. The latter genus is restricted to 8 American species.

As pointed out several times in recent studies of Atlanto-East Pacific stomato-
pods (Manning 1969:102; 1975:365; 1977:117), the 5 East Atlantic and South Af-
rican species assigned to the genus Meiosquilla Manning, 1968, differ from all
American species in having 5 rather than 4 spines on the dactylus of the raptorial
claw and in lacking elongate spines on the inner margin of the basal prolongation
of the uropod.

One of us (Manning 1962:507; 1969:103) suggested that some of the features
exhibited by species of Meiosquilla, including the unarmed anterolateral angles
of the carapace, the reduced carination of the carapace, thorax, and abdomen,
and the movable apices of the submedian teeth of the telson, are all characteristic
of squillid postlarvae. Thus, Meiosquilla sensu lato may have had a neotenic
origin. That the American and East Atlantic/South African species differ in 2
major features, number of teeth on the claw and ornamentation of the basal
prolongation of the uropod, suggests that these groups of species were derived
independently from different stocks and that they should not be assigned to the
same genus. The type-species of Meiosquilla is the West Atlantic Squilla quad-
ridens Bigelow, 1893, by original designation by Manning (1968:127). We recog-
nize here a new genus for the 5 East Atlantic and South African species formerly
assigned to Meiosquilla, which 1s here restricted to the 8 American species now
assigned to it (Manning 1969, 1972).

We thank C. W. Hart, Jr., Smithsonian Institution, and David K. Camp, Florida
Department of Natural Resources, for their comments on the manuscript.

Rissoides, new genus

Definition. —Body smooth, compact, size small to moderate, total length to 8.5
cm. Eye large, cornea bilobed, noticeably broader than stalk; ocular scales sep-
arate. Carapace smooth, narrowed anteriorly, anterolateral angles unarmed; ca-
rinae reduced, median and intermediates absent, at most reflected marginals and
posterior part of each lateral carina present; cervical groove indistinct; posterior
median margin evenly concave, posterolateral margins broadly rounded. Exposed
thoracic somites with, at most, intermediate carinae, submedians absent; lateral
process of fifth thoracic somite an inconspicuous diagonal or flattened lobe, a
ventral spine present on each side; lateral processes of next 2 somites evenly
rounded, not bilobed; ventral keel of eighth thoracic somite well developed. 4
epipods present. Mandibular palp absent. Dactylus of raptorial claw with 5 teeth,
outer margin evenly convex; propodus with outer part of upper (opposable) mar-

VOLUME 95, NUMBER 2 B55

gin evenly pectinate, with 3 proximal movable teeth, middle smallest; dorsal ridge
of carpus indistinct; ischiomeral articulation terminal. Endopods of walking legs
slender, elongate. Abdomen with anterior 5 somites lacking submedian carinae;
intermediates, laterals, and marginals present, usually unarmed anterior to fifth
somite; sixth abdominal somite with armed submedian, intermediate, and lateral
carinae. Telson broad, median carina present, supplemental dorsal carinae ab-
sent; 3 pairs of marginal teeth present, submedians with movable apices; pre-
lateral lobes absent; postanal keel, if present, low. Basal prolongation of uropod
with inner spine the longer, lacking elongate spines but with low tubercles or
short denticles on inner. margin; low, rounded lobe present on outer margin of
inner spine.

Type-species.—Squilla desmaresti Risso, 1816.

Etymology.—We consider it appropriate to dedicate this genus to A. Risso, the
nineteenth century carcinologist who named the type-species. The gender is mas-
culine.

Included species.—Five: Rissoides desmaresti (Risso, 1816); R. pallidus (Gies-
brecht, 1910); R. africanus (Manning, 1974); R. calypso (Manning, 1974); and R.
barnardi (Manning, 1975). Original citations for all of these species are given in
Manning (1977). .

Literature Cited

Manning, Raymond B. 1962. Alima hyalina Leach, the pelagic larva of the stomatopod crustacean
Squilla alba Bigelow.—Bulletin of Marine Science of the Gulf and Caribbean 12(3):496-507,
figures 1-4.

. 1968. A revision of the family Squillidae (Crustacea, Stomatopoda), with the description of
eight new genera.—Bulletin of Marine Science 18(1):105—142, figures 1-10.

. 1969. Stomatopod Crustacea of the western Atlantic.—Studies in Tropical Oceanography,
Miami 8:vii + 380, figures 1-91.

. 1972. Stomatopod Crustacea. Eastern Pacific Expeditions of the New York Zoological So-
ciety.—Zoologica, New York 56:95-113, figures 1-3.

. 1975. A new species of Meiosquilla (Crustacea, Stomatopoda) from South Africa.—Annals
of the South African Museum 67(9):363—366, figure 1.

. 1977. A monograph of the West African stomatopod Crustacea.—Atlantide Report 12:25—
181, figures 1-57.

(RBM) Department of Invertebrate Zoology (Crustacea), National Museum of
Natural History, Smithsonian Institution, Washington, D.C. 20560; (ChL) De-
partment of Zoology, Tel-Aviv University, Ramat-Aviv, Tel-Aviv, Israel.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 354-357

NEW RECORDS OF PINNOTHERID CRABS
FROM THE GULF OF CALIFORNIA
(BRACHYURA: PINNOTHERIDAE)

Mary K. Wicksten

Abstract.—Pinnotheres margarita is illustrated for the first time. The range is
extended northward in the Gulf of California. Pinnixa valerii is reported for the
first time from western Mexico. Illustrations are given of P. valerii and the closely
related species P. richardsoni.

Pinnotherid crabs are commensals with other invertebrates: annelids, echiu-
roids, mollusks, echinoderms, and ascidians. The species of the western coast of
Mexico are poorly known, often reported from single specimens of a given species
collected incidentally with other invertebrates.

Recent collecting by Alex Kerstitch, Ernest Iverson, and Michel Hendrickx
has resulted in the finding of many pinnotherid crabs. Among the specimens are
two species not recorded for more than 45 years—Pinnotheres margarita and
Pinnixa valerii. The new specimens have been deposited at the Allan Hancock
Foundation, University of Southern California; and the Estacion Mazatlan.

Pinnotheres margarita Smith
Fig. 1

Pinnotheres margarita Smith, 1869:245.—Smith, 1870:166.—Holmes, 1894:564.—
Rathbun, 1918:91-—93.—Glassell, 1934:301.—Schmitt, McCain, and Davidson,
1973:56—5S7.

Previous records.—Bay of Panama (type-locality), in pearl oyster (Pinctada
mazatlanica (Hanley), as Margaritiphora fimbriata).—La Paz, Muleje Bay (Gulf
of California, Mexico) (Rathbun 1918).

Material examined.—Guaymas, Sonora (27°54'N, 110°53’W), 10 m, rocky bot-
tom, commensal in Pinctada mazatlanica, 28 June 1981, A. Kerstitch, 1 female.—
Punta Chivato, Baja California (27°08’N, 111°54’W), 20 m among rock, sand and
algae, commensal in P. mazatlanica, 28 June 1980, A. Kerstitch, female, oviger-
ous.—SE side Bahia Concepcion, Baja California (26°43’N, 111°53’W), 18 Aug.
1980, E. Iverson, taken while snorkeling, female.—Isla Carmen, Gulf of Califor-
nia (25°58’N, ee 25 m, EnGHE rubble and small rocks, 10 July 1980, A.
Kerstitch, female.—**Panama,”’ 1866, F. H. Bradley, female.

Measurements in millimeters. —Carapace widths 11.5, 14.1, 10.7, 8.5, and 3.3
respectively; carapace lengths (in same order) 10.6, 12.3, 10.6, 8.2, and 3.1.

Remarks.—Pinnotheres margarita is one of the largest pinnotherid crabs in the
Gulf of California. However, the lack of illustrations has made identification of
the species difficult. The holotype, reported to have been deposited at the Pea-
body Museum of Natural History of Yale University, could not be located there
or at the National Museum of Natural History, Smithsonian Institution.

VOLUME 95, NUMBER 2 S55)

C

ys
ms
) I gfe
ss pu M
wil
A!
Sy
ih

Ss

_

—=55

—
SS

Fig. 1. Pinnotheres margarita, female from Punta Chivato, Baja California: a, Dorsal view; b,
Third maxilliped; c, Frontal region; d, Chela; e, Abdomen.

A small female, questionably identified as P. margarita, was found at the Pea-
body Museum. I compared this juvenile with the females recently collected. The
small crab has much the same shape of carapace, proportions of chelae and
walking legs, and outline of third maxilliped as the larger females. It also agrees
with the descriptions of Smith (1869, 1870) and Rathbun (1918).

Although type material and previous illustrations are lacking, I believe that the
females from the Gulf of California are indeed P. margarita. Smith and Rathbun
both mentioned the inequality in lengths of the walking legs on the right and left
sides in mature females. As described, the females are dull brown (‘‘like a uniform
coat of mud’’), with short pubescence. The carapace is uneven, with a protuber-
ant cardiac region and marked sutures. Reported size of the holotype (11.8 mm
in length of the carapace, 13.4 mm wide) agrees with measurements for the series
of crabs from the recent collections. Finally, the host (Pinctada mazatlanica) is
the same. All of the crabs agree with the previous descriptions.

356 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. a, Pinnixa valerii, male from Estero El Verde, Sinaloa; b, Pinnixa richardsoni, male
holotype from Balboa, Canal Zone, Panama.

Pinnixa valerii Rathbun
Fig. 2a

Pinnixa valerii Rathbun, 1931:262—263, figs. 1-2.—Schmitt, McCain, and David-
son, 1973:124.

Previous record.—Isla San Lucas, west coast of Costa Rica (type-locality), 15
Jan. 1930, 1 male holotype, 1 female paratype (Rathbun 1931).

Material examined.—Estero El Verde, Sinaloa (23°25’'30"N, 106°33’30"W), 1
m, among Ruppia sp., dredged, 11 Dec. 1979, M. E. Hendrickx and party.

Measurements in millimeters.—Carapace width 9.3, carapace length 5.8 (spec-
imen from Allan Hancock Foundation).

Comments .—Pinnixa valerii is related closely to Pinnixa richardsoni Glassell,
1936 (type-locality Balboa, Canal Zone, Panama). Both have laterally compressed
chelae with thick tufts of setae along the ridges of the palm and fingers. The
dactyls of the first pereopods appear twisted in dorsal aspect. Pinnixa valerii has
been illustrated previously only by photographs (Rathbun 1931, figs. 1 and 2).
Only the third maxilliped of P. richardsoni has been figured (Glassell 1936, pl.
ZY fies 3):

VOLUME 95, NUMBER 2 357

I compared the types of the two species (P. valerii from the U.S. National
Museum of Natural History and P. richardsoni from the San Diego Museum of
Natural History). In P. richardsoni, the merus of the third pereopod is 1.9x as
long as wide; in P. valerii, it is 2.7< as long as wide. The carpus of the third
pereopod in P. richardsoni bears a tubercle, absent in P. valerii. The first three
segments of the abdomen of P. richardsoni are fused, not articulated as in P.
valerii. In general, the legs of P. richardsoni (Fig. 2b) are much stouter than those
of P. valerii.

Some of the features given by Glassell in distinguishing between the two species
vary from animal to animal. Tomentum occurs on the outer surface of the hands
and carpus of the chelipeds of both species, differing merely in degree instead of
presence or absence. The outer distal margin of the third maxilliped appears
somewhat arched in two specimens of P. valerii, not angular. Both species have
the same twisted shape of the propodus and dacty] of the first pereopods.

So far, hosts for neither of these species have been recorded. Perhaps host
specificity will help in identification of these closely related species.

Pinnixa valerii was collected in an estuary at a salinity of 22%o. Future collec-
tors might seek this species in back bays, swamps, or estuaries.

Acknowledgments

I thank Michel Hendrickx, Estacion Mazatlan, for collecting specimens and
contributing toward preparation of the illustrations. Alex Kerstitch, University
of Arizona; Ernest Iverson, University of Southern California; and James
McLean, Los Angeles County Museum of Natural History, provided specimens
and information on hosts. John Garth, University of Southern California, gave
helpful criticism of the manuscript. The illustrations were prepared by Matilde
Méndez (Fig. 1), Frances Runyan (Fig. 2a), and Paula Walker (Fig. 2b). This
work was sponsored in part by a grant from Texas A&M University.

Literature Cited

Glassell, S. A. 1934. Affinities of the brachyuran fauna of the Gulf of California.—Journal of the

Washington Academy of Sciences 24:296-302.

. 1936. New porcellanids and pinnotherids from tropical North American waters.—Transac-

tions of the San Diego Society of Natural History 8:277-304.

Holmes, S. J. 1894. Notes on west American Crustacea.—Proceedings of the California Academy of
Sciences 4, series 2:563—588.

Rathbun, M. J. 1918. The grapsoid crabs of America.—Bulletin of the United States National Museum

97: 1-461.

. 1931. A new species of pinnotherid crab from Costa Rica.—Journal of the Washington

Academy of Sciences 21:262-263.

Schmitt, W. L., J. C. McCain, and E. S. Davidson. 1973. Decapoda I, Brachyura I. Fam. Pinno-
theridae. Jn Gruner, H.-E., and L. B. Holthuis, eds. Crustaceorum Catalogus. Den Haag, W.
Junk B.V. 160 pp.

Smith, S. I. 1869. Pinnotheres margarita Smith, sp. nov. In Verrill, A. E. On the parasitic habits of

Crustacea.—American Naturalist 3:245.

. 1870. Notes on American Crustacea. No. 1. Ocypodoidea.—Transactions of the Connecticut

Academy of Sciences 2:113-—176.

Department of Biology, Texas A&M University, College Station, Texas 77843.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 358-370

STUDIES OF NEOTROPICAL CADDISFLIES, XXXII:
THE IMMATURE STAGES OF MACRONEMA VARITPENNE
FLINT & BUENO, WITH THE DIVISION OF MACRONEMA

BY THE RESURRECTION OF MACROSTEMUM
(TRICHOPTERA: HYDROPSYCHIDAE)

Oliver S. Flint, Jr. and Joaquin Bueno-Soria

Abstract.—The larva and pupa of Macronema variipenne are described and
figured, and notes on their natural history given. A neotype is established for M.
agnathum Miller and it and Leptonema apicale Navas are synonymized with M.
bicolor Ulmer, and M. pullatum Navas is synonymized with M. trigramma Na-
vas. After considering all characteristics of variipenne and related species, Mac-
ronema is divided into two genera: Macronema Pictet, with M. lineatum Pictet
as type, is exclusively Neotropical; and Macrostemum Kolenati, with Macro-
nema hyalinum Pictet as type, contains not only Neotropical species, but all the
species presently assigned to Macronema in the Nearctic, Oriental, Ethiopian
and Australasian Regions.

The genus Macronema, as it has been recognized, is a large genus containing
somewhat over 100 described species. The genus is most diverse in the tropics
of the New World, Australia, Asia, and Africa, with a few species in North
America and northeastern Asia. It is notably absent from Europe and most of
northern Asia.

Although many adults have been described, very few larvae have been carefully
associated with their adult stage, and most of these are from the northern hemi-
sphere. In spite of the lack of associated material, many larvae of the Macrone-
matinae have been described, often incompletely, and then attributed to some
genus or species purely on supposition. This has, as one can imagine, produced
an extremely confused mess. Ulmer (1957, pp. 332-335) made a heroic attempt
to straighten out this confusion, and, in general, succeeded.

He united, under the name of Centromacronema auripenne (Ramb.), a series
of descriptions of a very distinctive larval and pupal type from South America.
This form was first made known through a series of works by Muller (1880, 1881,
1921 and variously referred to as Macronema, Macronema agnathum, Macro-
nema Species, third species, species C, and Macrosternum), and later fully sup-
plemented by the works of Thienemann (1905 as Macronema), Marlier (1964 as
Centromacronema), Roback (1966 as Hydropsychidae sp. 2), and Botosaneanu
& Sykora (1973 as Centromacronema). This larval type has often appeared in our
collections in small numbers, but under circumstances that never permitted a firm
association of stages. Finally we realized that these larvae were generally found
on plants, often in attached, free-floating roots or leaves, or in and around the
roots of emergent plants. With this information, Bueno was able to bring back to
the laboratory and rear in an aquarium several adults from a series of these larvae.
The adults that emerged turned out to be our recently described species, Mac-

VOLUME 95, NUMBER 2 So)

Figs. 1-4. Macronema bicolor, male: 1, Genitalia, lateral; 2, Same, dorsal; 3, Tip of aedeagus,
dorsal; 4, Same, lateral.

ronema variipenne, not a species of Centromacronema which also flies on the
same stream.

We take this opportunity to describe and figure the larvae and pupae of vari-
ipenne, select and figure a male neotype of M. agnathum Miller, synonymizing
it and Leptonema apicale Navas with M. bicolor Ulmer, and finally to discuss
the generic implications of this evidence, restricting Macronema to a group of
exclusively Neotropical species and resurrecting Macrostemum for another group
of worldwide distribution.

Macronema bicolor Ulmer
Figs. 1-6

Macronema bicolor Ulmer, 1905a:75. Fischer, 1963:178. Flint, 1966:6.

Macronema agnathum Miller, 1921:530. Ulmer, 1957:338. Fischer, 1963:177.
(New Synonymy)

Leptonema apicale Navas, 1927:40. (New Synonymy)

Because the identity of the larvae described by Miiller under the name Mac-
ronema agnathum has been unknown, and this uncertainty has been in large part
responsible for the confusion referred to above, we have made a concerted effort
to locate material from Muller. We have located some material in the Museum

360 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 5-6. Macronema bicolor: 5, Wings of lectotype; 6, Wings of neotype of M. agnathum.

VOLUME 95, NUMBER 2 361

2 ae gee = = — :
Soot rat se gy ws ~
Figs. 7-9. Habitats: 7, Rio Palma, above La Palma, Veracruz, Mexico; 8, Marginal vegetation; 9,
Aroid plant removed, in whose roots larvae were found.

of Comparative Zoology, Harvard University, and the Natural History Museum,
Vienna. In this latter collection are an adult male and female and two pupal skins
in excellent condition, all labeled in Miiller’s hand ‘‘Macronema III,’’ and the
two adults also **19/8 87.’’ Because the two adults agree with the brief diagnosis,
and the pupae are in total agreement with the more extensive description and
figures in Miller (1921), and are from him personally, we believe this material to
authentically represent Miiller’s concept of agnathum and are therefore labelling
the male **Neotype.”’

The neotype is the same species as M. bicolor Ulmer with which it is here
synonymized. Ulmer (1957, p. 341) also noted (courtesy of Mr. D. E. Kimmins)
that the British Museum (Natural History) had examples labeled “‘III’’ and that
these too were bicolor, but then decided that III referred to M. chloraemus
Miller, not agnathum. It appears that he was confused by the erratic usage of
letters, numbers, and names by Miller.

The type male of Leptonema apicale Navas located in the Deutsches Ento-
mologisches Institut, Akademie der Landwirtschaftswissenschaften, was bor-
rowed (through the kindness of Dr. G. Petersen) and studied. It is a rather faded
example that matches bicolor in all respects, with which it is herewith synony-
mized.

The figures of the male genitalia here presented (Figs. 1-4) were prepared from
the neotype of agnathum, as is the photograph of the wings (Fig. 6). The neotype

362 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

11 12

Figs. 10-12. Macronema variipenne, larva: 10, Lateral; 11, Head, dorsal; 12, Thorax, dorsal.

is in excellent, unrubbed condition, whereas the types of bicolor and apicale are
rubbed and appear rather different in coloration (Fig. 5).

Macronema variipenne Flint & Bueno
Figs. 7-24

Macronema variipenne Flint & Bueno, 1979:528—532.

VOLUME 95, NUMBER 2 363

\S
La wa
S\ Y
AY

Figs. 13-19. Macronema variipenne, larva: 13, Maxillolabium and anterior region of genae, ven-
tral; 14, Mandibles, ventral; 15, Labrum, dorsal; 16, Forepleuron and coxa, lateral; 17, Foreleg,
lateral, with 2 margin hairs of femur enlarged; 18, Midleg, lateral; 19, Prosternite, ventral.

This species is widespread through eastern and southern Mexico, south
throughout Central and South America at least as far as Ecuador. We have taken
adults commonly at lights at night as well as by net during the day.

Larva.—Length to 21 mm, width 2.5 mm. Sclerites mostly golden-yellow with
darker brown markings.

Head: As wide as long. Color brown dorsomesally, golden-yellow laterally,
posteriorly, and ventrally but becoming darkened ventromesally; frontoclypeus
with a pair of pale spots laterally. Frontoclypeus with anterior margin concave,
with prominent knobs anterolaterally; surface with only a few setae laterally.
Genae with surface smooth, beset with short, bladelike setae in a broad band

364 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

23

Figs. 20-23. Macronema variipenne: 20, Larval anal proleg, lateral; 21, Pupal head, anteroventral;
22, Pupal hook plates, with segment number and anterior or posterior positions indicated, plates 5A
to 8A shown in dorsal aspect to left, in lateral aspect to right; 23, Apex of pupal abdomen, dorsal.

VOLUME 95, NUMBER 2 365

Meso Meta | 2 3 A 5 6 7! 8

Fig. 24. Macronema variipenne, schematic gill diagram of the lateral aspect of the meso- and
metathorax and first 8 abdominal segments. S = a single long central stalk with numerous lateral
filaments. D = 2 S-type gills arising from a common base.

laterally and posteriorly (but not on pale area posteriorly); with scattered long
setae. Genae without stridulatory grooves ventrally; ventral ecdysal line lacking
on one side anteriorly, thus anterior ventral apotome is only half delimited. La-
brum broad, rounded anterolaterally; anterior half hairy; no anterolateral brushes.
Mandibles short, broad, lacking mesal teeth and brushes; apicomesal region strik-
ingly black internally. Submentum with anterior margin convex, with a few setae
at anterolateral angles.

Thorax: Pronotum brown on anterior half, yellow on posterior half; meso- and
‘metanota golden-yellow with fuscous lateral and posterior margins. Pronotum
with bladelike setae and long hairs on anterior half, posterior half smooth; meso-
notum with a few shorter bladelike setae mostly anterolaterally and many long
and short hairs; metanotum with only long and short hairs. Prosternite almost
crescentic with an anteromesal projection; with a small, triangular sclerite pos-
teromesally. Meso- and metasterna unornamented except for a few short setae.
Foretrochantin tapering to a blunt point, with | bladelike seta; femur with a
ventral row of 8 palmately divided setae; all segments with numerous hairs. Mid-
and hindlegs very similar in structure: coxa, femur and trochanter very elongate;
trochanter and segments distad, narrow and terete, with rows of short, peglike
setae and long and short hairs; tarsus with a single apicodorsal bladelike seta;
claw almost straight, with basal setae almost as long as claw. No thoracic gills.

Abdomen: Gills only laterally as shown in Fig. 24. Lateral line lacking. Integ-
ument with long and short hairs dorsally, smooth ventrally. Sternum 8 with scat-
tered hairs including 2 pairs of long setae arising from a pair of small, indistinct
sclerites. Sternum 9 very hairy, bearing a pair of large, pale, hairy sclerites which
also bear a few short, bladelike setae anteriorly; a small narrow sclerite laterad.
With 4 long, slender anal gills. Anal prolegs extremely long and slender; ventro-
basal plate (attached to the abdomen) bearing a pair of longitudinal dark stripes;
legs very long, slender, and hairy with a thin section of cuticle at midlength at
which point the legs bend; apical brush reduced to 1 or 2 hairs; claw long, slender,
curved.

Pupa.—Length to 10 mm, width 3 mm.

Head: Labrum semicircular, with rounded basolateral lobes; anterior third
densely hairy, basolateral angles with 5-6 short setae, basolateral lobes with 5-
6 setae. Mandibles membranous, triangular, anterior margin broadly truncate.

366 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Face with a few scattered setae; anterior tentorial pits well marked. Basal anten-
nal segment with a group of setae; antennae very long, curled into several broad
loops (not wrapped around the apex of the abdomen).

Thorax: Meso- and metanota with a few scattered setae and a cluster of setae
on the tegulae of mesonotum. Midlegs with tibiae and tarsi flattened and fringed.

Abdomen: Segments 3-7 with long, terete lateral processes. Gills laterally be-
tween processes and ventrally. Hook plates anteriorly on segments 3-8 (that of
4 very small, those on 5-8 very high); posteriorly on 3 and 4. Segments 3-8 with
a few pairs of setae posteriorly. Segments 8 and 9 with a dense fringe of black
hairs laterally. No apical appendages, only a cluster of black hairs.

Biology.—The larvae were found in the free-floating mats of roots, and in the
tangled roots and stems of emergent plants. The retreats in the root mats were
about 30 mm long by 7 mm wide, constructed of fragments of organic matter held
together rather loosely by silk and all attached to the rootlets. These retreats
were removed as carefully as possible and were preserved promptly, but the
larvae generally crawled away quickly. The retreats were later cut open carefully,
in an attempt to find some sort of capture net. Nothing of any type could be
found, and the retreats seemed to be only a simple tube. We believe that the
larvae construct no distinct net, either of the Macronema type or of the Hy-
dropsychinae type. Perhaps they may construct an irregular mesh of threads
outside the shelter, but even that seems unlikely.

Pupal shelters have also been found in the same root mats. These are approx-
imately the same size as the larval shelters, but are a little larger in diameter
where the pupa is, and are more firmly constructed. Inside the shelter of silk and
detritus, the larva constructs a perfect silken cocoon, free of the outer shelter
except near each end. The ends are simply a loose tangle of threads. None of the
pupae found in the wild was mature enough to permit species identification.

In order to determine the species of these larvae, Bueno, in the summer of
1978 collected a series of larvae in the Rio Palma (Fig. 7), above the village of La
Palma. This site is in the tropical wet forest of the coastal lowlands in the state
of Veracruz, about 25 km east of Catemaco at an elevation of 30 m above sea
level. The river is no more than 15 m wide with an average depth of about 14 m,
the water is clear, of good chemical and physical condition, and the bottom is
sandy with much gravel and stone. At the sides there are often backwaters with
emergent plants, and the marginal trees and vines often put into the water roots
which greatly proliferate, frequently forming free-floating mats. The large leaves
of the marginal trees and shrubs form a favored resting place for the adults during
the day.

The larvae, mostly in the 4th and Sth instars, were collected from the roots and
stems of the emergent plants (Figs. 8-9), and then brought back alive to the
laboratory in Mexico City, a distance of some 550 km. There they were kept
alive in an aquarium in water with sand, sticks and debris all from the Rio Palma.
The aquarium was of about two liters capacity, and air was supplied by an air
pump which at the same time caused the water to circulate in the aquarium. A
supply of ‘““Tetramin,’’ a tropical fish food, was circulated with the water every
day to provide food for the larvae. When most of the water had evaporated, it
was replaced with water directly from the tap.

The larvae mostly disappeared into the bottom sand and debris immediately.

VOLUME 95, NUMBER 2 367

However, a few constructed their retreats on the glass and could be observed in
their tunnel. These larvae spent most of their time simply resting in their tube.
Rarely they would crawl part way out and appear to be, perhaps, feeding on
debris on the retreat. When the water level became low, the current diminished,
and they spent much of their time making undulatory movements with the ab-
domen. After a while these larvae disappeared, and in about 20-30 days adults,
which were M. variipenne, emerged.

We have examined the gut contents of several field preserved larvae. The guts
were filled with rather uniform, roughly cubical, bits of plant tissue about 0.2 mm
long on a side.

In summary, it is our belief that the larvae: construct only a rather flimsy,
tubular larval retreat without any type of capture net; construct the retreat on
plant structures in marginal situations or even buried in silt or sand in backwaters;
that they are very tolerant of low water velocities and oxygen concentrations;
and that they are “‘shredders,’’ biting-out pieces of plant material. The pupae are
sheltered in a slightly enlarged and strengthened part of the larval retreat; the
pupa is surrounded by a silken cocoon only attached to the outer shelter near the
ends; and that the ends of the cocoon are not firmly closed, but loosely plugged
by an irregular tangle of silk.

Generic Considerations

Now, with the first unequivocal association of this form of larva with a species
of the genus Macronema, not Centromacronema as it was believed to be (Ulmer
1957; Marlier 1964), the entire question of generic relationships is thrown open.

The descriptions of the immature stages of a number of species of ‘‘Macro-
nema’ have been in the literature for many years: carolina Bks. (Ross, 1944),
zebratum Hag. (Ross, 1944), transversum (Walk.) (Ross, 1944), radiatum McL.
(Lepneva, 1964), capense Walk. (Scott, in press), u/meri Bks. (Marlier, 1964, as
siolii), and others described but not associated with any adult (as listed in Ulmer
1957, pp. 344-345). All these descriptions are very similar and are summarized
in Table 1 (under Macrostemum) where the contrasting characteristics of vari-
ipenne are also given (under Macronema). 1 do not believe any trichopterist
would consider that such totally discordant characteristics could belong to dif-
ferent species in one genus, especially so if they considered the characteristics
of other hydropsychid genera (as shown in Wiggins 1977, for example). No won-
der Ulmer (1957, p. 339) concluded that the variipenne type larva could not be
a Macronema, and assigned it to the closely related genus Centromacronema.

Unfortunately, we still do not have any larvae definitely correlated with Centro-
macronema, nor with some of the more distinctive species of Macronema from
eastern Brazil. Such a lack, however, does not affect the problems within Mac-
ronema, but does prevent a complete solution incorporating all the closely related
genera. Recognizing the possibility of the future synonymy of Centromacronema
with Macronema, and the possibility of having to carve more genera from Mac-
ronema, we will consider the situation within Macronema, as these aforemen-
tioned possibilites should not affect the stability of the names in consideration.

Ulmer was the first to give notice in a publication (1907) of two distinct groups
of species in Macronema. The first group of exclusively South American species

368

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Comparisons of certain characters of the larva and pupa of Macronema and Macroste-

mum.
Macronema Macrostemum
Larva
Labrum no brushes anterolateral brushes
Mandibles short, no teeth elongate, toothed
Gena, dorsally unmodified with a carina around central

Gena, ventrally
Ventral ecdysal line

Coxa
Foretibia and tarsus

Mid- and hind tibiae and
tarsi

Thoracic gills
Abdominal gills

Anal prolegs
Retreat

Net

Food

Pupa

Mandibles

Labrum

Antennae

Posterior hook plates
Apical appendages
Inner cocoon
Cocoon ends

no stridulatory grooves
lacking on | side of pregula

long

scattered hairs, ventral row of
palmate setae

disproportionately short in
relation to femur; with rows
of short, pointed setae

lacking
lateral line only

very long, jointed in middle
tubular
none

shredders, biting pieces from
plant matter

membranous

hairy on anterior third
looped only

on segments 3 and 4

lacking

free of shelter except at ends
loosely plugged

area
with stridulatory grooves

separating pregula on both
sides

short
with a large brush of hairs

typical proportions; with
ventral rows of feathered
setae

present

dorsolateral, lateral, and
ventrolateral rows

normal, not jointed
branched
very regular, of fine mesh

collectors, gathering suspended
fine particulate matter

sclerotized

few scattered setae
coiled around abdomen
on 3 only

present

united to shelter
closed

(in which he included lineatum, argentilineatum, parvum, fulvum, bicolor, and
percitans) he characterized by a minute discoidal cell, a very large median cell,
a distinct, extra, costal crossvein, and the subcostal vein ending on the costa. The
other group, consisting of all the other species in the genus, possessed a normal
or large discoidal cell, a smaller median cell, no costal crossveins, and the sub-
costa was united to R, just before the wing margin. However, Miiller had noticed
as early as 1885 (but not published until 1921) a division of the genus based on
the presence or absence of the apical tibial spurs on the foreleg and structure of
the pupa (as in Table 1). Kolenati (1859) had proposed a new genus Macrostemum
for M. hyalinum Pict. and M. auripenne Ramb. based on the absence of such
spurs in the latter two species. M. hyalinum does have a pair of tibial spurs, but
they are very small and easily overlooked in comparison to the large spurs of
those species he placed in Macronema (lineatum although included, was appar-

VOLUME 95, NUMBER 2 369

ently unknown to him). Ulmer in 1957 selected hyalinum as the type of Macros-
temum (but considered it and Macronema synonymous), and did correctly at-
tribute the species with the two small spurs to it and the ones lacking spurs to
Macronema. In 1979 we independently recognized the same two groups on the
basis of coloration and male genitalia.

Now with the knowledge of the larval and pupal morphology and ecology, it
is obvious that these differences between the two groups of species in the adult
stage are much more fundamental. Therefore, we propose to recognize two gen-
era. Macronema Pictet (1836) has as its type M. lineatum Pictet (monobasic) and
in addition contains: amazonense Flint, argentilineatum Ulm., bicolor Ulm.,
bifidum Flint, burmeisteri Bks., chalybeoides Ulm., exophthalmum Flint, fragile
Bks., fraternum Bks., fulvum Ulm., gundlachi Bks., hageni Bks., immaculatum
Mos., lachlani Bks., luteipenne Flint & Bueno, matthewsi Flint, muelleri Bks.,
paliferum Flint, parvum Ulm., pennyi Flint, percitans Walk., pertyi Bks., picteli
Bks., reinburgi Nav., and variipenne Flint & Bueno. All species are limited to
the Neotropical Realm.

The salient characteristics distinguishing the adults of this genus from Mac-
rostemum are: lack of tibial spurs on the forelegs; forewing with Sc ending on
the costa; forewing color due mostly to scales, basal %4 usually being green,
bounded outwardly by a variably colored area and with costal cell filled with
silvery scales; male with claspers undivided or nearly so, tenth tergum shorter
and broader, often with lobes and processes, tip of the aedeagus also often bears
lobes and processes.

The larval and pupal characteristics are outlined in Table 1.

The second genus is Macrostemum Kolenati (1859), with the type Macronema
hyalinum Pictet (selected by Ulmer 1957). In addition to the type the genus con-
tains the following species in the New World: arcuatum Erich., braueri Bks.,
carolina Bks., digramma McL., erichsoni Bks., maculatum Perty, negrense
Flint, par Nav., ramosa Nav., santaeritae Ulm., subequale Bks., surinamense
Flint, transversum Walk., trigramma Nav. (pullatum Nav. is a NEW SYN-
ONYM), triste Nav., tuberosum Ulm., ulmeri Bks., and zebratum Hag. In ad-
dition, all species listed in the Fischer catalogs (1963, 1972) in Macronema from
Africa, Asia, and Australia are transferred to Macrostemum.

The adults differ from Macronema by a combination of the following charac-
teristics: foretibia with 1 or 2 small apical spurs; Sc of forewing united with R,
apically, or ending in a fork whose ventral arm is united to R, and is the stronger
of the two arms; forewings with color due primarily to strongly contrasting colors
of the membrane and the pattern is widespread over the wing; males with claspers
distinctly 2-segmented, the tenth tergum is elongate and rather simple, and the
tip of the aedeagus is generally bulbous without special structures.

The larval and pupal structures of this genus are also outlined in Table 1.

Acknowledgments

We express our appreciation to the Smithsonian Staff artists, Young T. Sohn
(Figs. 11, 17, 18, 20-23) and L. Michael Druckenbrod (Figs. 10, 12), for their
excellent illustrations. Drs. Richard C. Froeschner and Wayne N. Mathis pro-
vided much useful discussion and criticism of the manuscript. We thank Jorge
Padilla and other students of the U.N.A.M. for their valuable help in the field.

370 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Flint is most grateful to the Director of the Instituto de Biologia, Universidad
Nacional Autonoma de Mexico, Dr. Jose Sarukhan Kermes, for travel funds and
to the Secretary of the Smithsonian Institution, Dr. S. Dillon Ripley for Fluid
Research Funds, which together made possible this work in Mexico. A CONA-
CYT grant permitted Bueno to complete the study at the Smithsonian Institution.

Literature Cited

Botosaneanu, L., and J. Sykora. 1973. Sur quelques Trichopteéres (Insecta: Trichoptera) de Cuba.—
Résultats des expéditions biospéologiques cubano-roumaines a Cuba 1:379-407.

Fischer, F. C. J. 1963. Hydropsychidae, Arctopsychidae.—Trichopterorum Catalogus IV:1-172.

. 1972. Supplement to Vol. III and I1V.—Trichopterorum Catalogus XIII:1-172.

Flint, O. S., Jr. 1966. Studies of Neotropical Caddis Flies, II: Types of some species described by
Ulmer and Brauer.—Proceedings of the United States National Museum 120(3559):1-21.

Flint, O. S., Jr., and J. Bueno-Soria. 1979. Studies of Neotropical Caddisflies, XXIV. The genus
Macronema in Mesoamerica (Trichoptera: Hydropsychidae).—Proceedings of the Entomolog-
ical Society of Washington 81:522-—535.

Kolenati, F. A. 1859. Genera et Species Trichopterorum. Pars Altera.—Nouveaux Mémoires de la
Société Imperiale des Naturalistes de Moscou X VII(XI): 141-296.

Lepneva, S. G. 1964. Larvae and pupae of Annulipalpia.—Fauna of the U.S.S.R., Trichoptera
II(1): 1-560.

Marlier, G. 1964. Trichopteres de | Amazonie recueillis par le Professeur H. Sioli.—Institut Royal
des Sciences Naturelles de Belgique, Mémoires(2nd ser.) 76:1—167.

Miller, F. 1880. Sobre as casas construidas pelas larvas de insectos Trichopteros da Provincia de

Santa Catharina.—Archivos do Museu Nacional (Rio de Janeiro) 3:99-134, 209-214.

. 1881. Ueber die von den Trichopterenlarven der Provinz der Santa Catharina verfertigten

Gehause.—Zeitschrift fiir wissenschaftliche Zoologie 35:47-87.

. 1921. Briefe und noch nicht veroffentlichte Abhandlungen aus dem Nachlass 1854-1897. Vol.

2:xvil + 667, 239 figs. In A. Moller, ed., Fritz Miiller: Werke, Briefe und Leben. G. Fischer,

Jena. 3 vols.

Navas, R. P. L. 1927. Neuropteren, Megalopteren, Plecopteren und Trichopteren aus dem Deutsch.
Entomolog. Institut (Berlin-Dahlem).—Entomologische Mitteilungen 16:37-43.

Pictet, F. J. 1836. Description de quelques nouvelles especes de Névropteres, du Musée de Genéve.—
Memoires de la Société de Physique et d’ Histoire Naturelle de Genéve 7:399-404.

Roback, S. S. 1966. The Catherwood Foundation Peruvian-Amazon Expedition, XI-The Trichoptera
larvae and pupae.—Proceedings of the Academy of Natural Sciences of Philadelphia 14:235-
303.

Ross, H. H. 1944. The Caddis Flies, or Trichoptera, of Illinois.—Illinois Natural History Survey,
Bulletin 23(1): 1-326.

Scott, K. M. F. In prep. On the Hydropsychidae (Trichoptera) of Southern Africa, with keys to the
African Genera of imagos, larvae and pupae and species lists.—Annals of the Cape Provincial
Museums (Natural History).

Thienemann, A. 1905. Trichopterenstudien.—Zeitschnft fiir wissenschaftliche Insektenbiologie
1:285—291 .

Ulmer, G. 1905. Zur Kenntnis aussereuropdischer Trichopteren.—Stettiner Entomologische Zeitung

66: 1-119.

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(OSF) Department of Entomolgy, Smithsonian Institution, Washington, D.C.
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PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 371-376

NOTES ON DISTRIBUTION OF SOME LATIN AMERICAN
COTTON-STAINERS (DYSDERCUS: PYRRHOCORIDAE:
HEMIPTERA) AND REMARKS ON THE BIOLOGY OF
DYSDERCUS URBAHNI SCHMIDT

Joachim Adis and Richard C. Froeschner

Abstract.—New geographical distribution data are given for 11 Latin American
species of Dysdercus, with new country and/or state records for 7 of them. Data
on life history and food sources are given for D. urbahni Schmidt in a riverine
inundation-forest in the Central Amazon, Brazil.

Introduction

Extensive taxonomic and distributional data for Latin American species of
Dysdercus were given by Ballou (1906), Lima et al. (1962), Doesburg (1968), and
Beccari and Gerini (1970). Now additional geographic distribution can be given
based on specimens recently located or deposited in Museu Paraense Emilio
Goeldi (MPEG), Belem, Brazil; Instituto Nacional de Pesquisas da Amazonia
-(INPA), Manaus, Brazil; and the Smithsonian Institution, National Museum of
Natural History (NMNH), Washington, D.C., U.S.A.

The first significant biological data for Dysdercus urbahni Schmidt were gath-
ered during 1975-1976 ecological studies in a white water, inundation-forest
(‘‘Varzea’’) near Manaus, Brazil.

Notes on the Distribution of Some Latin American Dysdercus

Present new geographic records, some based on identifications by P. H. van
Doesburg in 1978, mostly complement the distribution data in Doesburg’s (1968)
revision of the New World species of Dysdercus.

Dysdercus mimus Group

1) Dysdercus imitator Blote

Previous records: Colombia, Venezuela, Ecuador, Peru, Bolivia (Doesburg
1968).

Present records: Brazil-Mato Grosso: Aripauna (5°07'S, 60°24’W), Reserva
Humboldt, II-1977, 1 6, W. L. Overal; Chapada dos Guimaraes (15°26’S,
55°45’W), 3-II-1961, 1 2, J. e B. Bechyné Coll. (all in MPEG).

2) Dysdercus mimus infuscatus Blote

Previous records: British Guiana, Suriname (Doesburg 1968).

Present records: Brazil-Amapa: Macapa, Praia do Araxa, 19-X-1978, 1 d/I @&,
M. F. Torres (MEPG).—Amazonas: Manaus 010 km 232 (=highway Manaus-
Itacoatiara km 232), 28-VI-77, 1 6, B. Mesearenhas (INPA).—Maranhao: Im-
peratriz (5°32'S, 47°29'W), 13-VI-1978, 1 ¢, F. F. Ramos; Santa Inés, 05-X-1978,
736/62 °¢,F. F. Ramos/P. Celso.—Mato Grosso: Aripuana, Reserva Humboldt,
IIl-1977, 1 6, W. L. Overal.—Para: Araguaia, Redencao, 25-XI-1978, 1 2, W.

372 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Franca, Belém, Mocambo, 6-V-1977, 1 6; 01-III-1977, 1 ¢, P. Waldir; 5-III-1977,
1 °,A. Y. Harada. Belém, Mocambo, 19-XII-1977, 2 2 2, C. Moreira; XII-1977,
1 2, C. Moreira; 02-I-1978, 1 2, S. A. Santos. Belém, Parque do Museu, X-1976,
2466/1 2, W. L. Overal. Belém, Utinga, 22-III-1977, 1 2, P. Tadeu; 22-ITI-1977,
1 6, M. F. Torres; 24-III-1977, 1 6, M. F. Torres. Rodovia Belém-Brasilia, km
90, F. S. Antonio, 14-II-1976, 1 6/9 2°, T. Pimentel; 14-II-1976, 3 dd/3 29,
M. F. Torres; 18-II-1976, 2 6d, M. F. Torres; 26-II-1976, 1 6, M. F. Torres.
Rodovia Belém-Brasilia, km 90, F. Jaboti, 13-II-1976, 2 64/2 2°, T. Pimentel;
13-II-1976, 1 2, M. F. Torres. Braganca (1°03’S, 46°46’W), 26-IV-1978, 1 ¢, W.
L. Overal; 26-V-1978, 3 64/4 22, F. F. Ramos/M. F. Torres. Bujaru (1°31’S,
48°01’W), 03-VIII-1978, 2 dd/2 22, P. Nolasco; 03-VII-1977, 1 2, P. Nolasco;
29-VI-1978, 1 ¢, P. Waidir/L. Braak (MPEG). Capitao Poco, 25-II-1978, 2 ¢6/
4 29, W. Franca/P. Tadeu; 26-II-1978, 1 6/5 2°, W. Franca/P. Tadeu; 27-II-
1978, 1 36/2 292, W. Franca; 28-I]-1978, 1 2, W. Franca. Ilha das Oncas
(=across Belém), 30-I-1977, 1 6, W. L. Overal. Maraba (5°21’S, 49°07'W), 3-
VII-1978, 1 o/1 29, W. Franca. Peixe-Boi (1°12’S, 47°18’W), 24-I-1964, 1 9, W.
Franca; 15-IV-1977, 1 2, P. Waldir; 16-IV-1977, 7 64/5 29, P. Waldir; 12-IV-
1977, 1 36 P. Waldir; 17-IV-1977, 1 2, W. L. Overal; Peixe-Boi, 24-I-1964, 1 ¢,
W. Franca; 22-IV-1978, 1 6, M. F. Torres; 23-IV-1978, 1 3/1 2, M. F. Torres;
24-IV-1978, 1 6, F. F. Ramos; 26-IV-1978, 1 2°, F. F. Ramos; 27-IV-1978, 1 @,
N. de Souza. S. J. Pirabas, 27-VII-1978, 1 6/1 2, M. F. Torres. Santa Isabel
(1°41’'S, 50°11’W), 29-I-1964, 1 3/2 2992, W. Franca. Tucurui (3°42'S, 49°27’ W),
7-VII-1978, 1 2, P. Tadeu; 8-VII-1978, 4 9°, R. B. Neto; 21-X-1978, 1 ¢, R. B.
Neto; 14-III-1979, 1 6, margem esquerda, W. L. Overal (all in MPEG).

These first Brazilian records for this subspecies extend its range coastwise and
far inland up the valley of the Amazon River.

Dysdercus flavolimbatus Group

3) Dysdercus ruficeps Perty

Previous records: Nicaragua; Costa Rica, Panama, Colombia, Ecuador, Peru,
Bolivia, Brazil: Amazon (6 records, only one with exact locality), Para (4 records,
one without exact locality), Rio de Janeiro (1 record; comp. Doesburg 1968,
Beccari & Gerini 1970).

Present records: Brazil-Para: Ananindeua (1°22’S, 48°23’W), III-1956, 1 2, C.
Carvalho; Belém, Fazenda Velha, 14-V-1977, 1 2, M. F. Torres; Belém, Utinga,
22-III-1977, 1 2, P. Tadeu; Benevides, VI-1964, 1 9, W. Franca; Benevides
(1°22’S, 48°15’W), VI-1964, 1 2, W. Franca; Bujaru, 01-VII-1977, 1 36, P. Waldir/
L. Braack; Mosqueiro (1°10’S, 48°28’W), 17-VII-1978, 3 63/1 2, M. F. Torres;
Santa Isabel (1°41’S, 50°11’W), 17-XII-1973, 3 66/3 2 2, B. Mascarenhas; 10-II-
1974, 5 64/2 22, José Queiroz; (all in MPEG).

4) Dysdercus flavolimbatus flavolimbatus Stal

Previous records: Mexico, Guatemala, Panama: Rio Changena, Boca del Toro,
2400 ft., 17-[X-1961, 1 36, G. B. Fairchild (comp. Doesburg 1968).

Present record: Panama-Chiriqui (Dist. Renacimiento): Santa Clara 4000 ft.
(8°31'S, 82°39’'W), 20-22-V-1977, 1 g6/2 22, B. C. Ratcliffe (INPA).

VOLUME 95, NUMBER 2 37/8

5) Dysdercus concinnus pehlkei Schmidt

Previous records: Costa Rica, Venezuela, Colombia, Panama: La Chorrera,
23-V-1944, K. E. Frick; Lino, 2 2 (comp. Doesburg 1968).

Present record: Panama-Colon: Santa Rita Ridge (9°20’N, 79°48’ W), 24-V-1977,
1 6, B. C. Ratcliffe (INPA).

Dysdercus maurus group

6) Dysdercus obscuratus flavipenuis Blote
Previous records: Panama, Colombia, Ecuador, Venezuela (Doesburg 1968).
Present record: Brazil-Mato Grosso: Chapada dos Guimaraes (15°26’S,
55°45’W), 5-II-1961, 1 2, J. e B. Bechyné (MPEG).

7) Dysdercus honestus Blote
Previous records: Peru, Colombia, Venezuela, Suriname, Paraguay, Bolivia,

Brazil: Amapa (1 record), Bahia, Espirito Santo, Goias (5 records), Mato Grosso
(1 record without exact locality), Minas Gerais, Para (3 records), Rio de Janeiro,
Sta. Catharina (comp. Doesburg 1968).

- Present records: Brazil-Goids: Pirineus, 2-II-1962, 1 ¢; Paraiso, 9-II-1962, 1
2, J. e B. Bechyné.—Maranhao: Buriticupu (4°13’S, 46°33’W), 30-IX-1978, 1
3, M. F. Torres.—Mato Grosso: Chapada dos Guimaraes (15°26’S, 55°45’W), 19-

mioivoleiece J. es BaiBechyne; 21-1-1961, 19, J: e B. Bechyneé; 5-11-1961, 1 95 J.

e B. Bechyné (all in MPEG). Rio (=Res.) Humboldt, Estrada do Porto (=Cidade

Humboldt; 5°07’S, 60°24’W), 1 6, L. Albuquerque, nr. 0240 (INPA); Rio Hum-

boldt, Estrada Dir. Aeroporto (=Cidade Humboldt), 26-I-1976, 1 ¢, (NMNH);

Fazenda Ric. Franco, 6-III-1961, 1 ¢6, J. e B. Bechyné; 7-III-1961, 1 3/1 @, J.

e B. Bechyné; 15-III-1961, 1 2, J.e B. Bechyné.—Para: Tucurui (2°53’S, 52°0’W)
14-IIJ-1979, 1 2, W. L. Overal, ‘‘margem esquerda’’ (both in MPEG).

Observation: The state record for Maranhao is the first record of D. honestus

in the northeast of Brazil (comp. Doesburg 1968, fig. 174).

8) Dysdercus ruficollis (Linnaeus)

Previous records: British Guiana, Suriname, Argentina, Paraguay, Brazil:
Amazonas, Bahia, Distrito Federal, Espirito Santo, Goias, Guanabara, Minas
Gerais, Parana, Rio de Janeiro, Rio Grande do Sul, Sao Paulo, Sta. Catharina
(comp. Doesburg 1968).

Present records: Brazil-Maranhao: Imperatriz (5°32'S, 47°29’W), 13-VI-1978,
7364/8 22, M. F. Torres/F. F. Ramos/R. B. Neto, (MPEG).—Rio de Janeiro:
Manguinhos, 30-V-1967, 1 6/2 22, L. P. Albuquerque; IV-1967, 1 3, L. P.
Albuquerque (INPA); Petropolis, 5-ITI-1962, 1 2, J. e B. Bechyné (MPEG).

Observation: The first state record for Maranhao indicates the presence of D.
ruficollis in the northeast of Brazil (comp. Doesburg 1968, fig. 210).

Dysdercus albofasciatus group

9) Dysdercus goyanus Doesburg

Previous record (comp. Doesburg 1968): Brazil-Goias: Jatahy (latai). Clermont
vend. (1 d, holotype).

Present record: Brazil-Mato Grosso: Cuiaba (15°35'S, 56°05’ W), 22-III-1961, 1
6, J. eB. Bechyné (MPEG).

374 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Dysdercus fernaldi group

10) Dysdercus urbahni Schmidt

Previous records: Brazil: Amazonas, Para (comp. Doesburg 1968).

Present records: Brazil-Amapa: Macapa, Pacoval, 07-XI-1978, 1 6, W. Franca
(MPEG).—Amazonas: Ilha de Curari (Rio Solimoes), 10-VII-1979, 1 S/1 2/6
nymphs (all in alcohol), J. Adis ‘‘em sementes de Pseudobombax munguba caidas
no solo’’; Iranduba, Rio Solimoes, 29-I[X-1962, 3 ¢ 36, J. Dellome, no. 2471; Estr.
AM 1 (=Manaus-Itacoatiara) km 180, 21-X-1965, 1 6, F. Antonio; Estr. AM 1
km 244, 11-I-1977, 1 ¢, B. C. Ratcliffe; Urucury, Codajas, XII-1963, 1 6, Ed-
uardo, no. 2552.—Mato Grosso: Aripuana (=Cidade Humboldt; (5°07’S,
60°24’W), Estrada do Porto, 1 6, L. P. Albuquerque/E. Rufino, no. 0180 (all in
INPA).

Observation: The state record for Mato Grosso indicates the presence of D.
urbahni in the southern Amazon region. (comp. Doesburg 1968, fig. 242). For
biology notes see p. 9.

11) Dysdercus fernaldi fuscofasciatus Blote

Previous records: Venezuela, British Guiana, Suriname, French Guiana, Bra-
zil: Amazonas, Para (1 record without exact locality), Sao Paulo (comp. Doesburg
1968).

Present records: Brazil-Amazonas: Urucury, Codajas, (6°46’S, 64°35’W), XII-
1963, 5 dd/5 22, Eduardo, no. 2551 (INPA).—Para: Ilha das Oncas (=across
Belém), 7-II-1977, 1 6, W. L. Overal; 30-II-1977, 1 2, W. L. Overal (MPEG).

Notes on the Biology of Dysdercus urbahni Schmidt

The following information comprises the first biological observations recorded
for Dysdercus urbahni. The species has been collected mainly along rivers (black
water; e.g. Rio Negro; white water: e.g. Rio Solimoes) and on ‘‘terra firme’’ (e.g.
Reserva Ducke, Manaus) in the Amazon area. The observations were made dur-
ing ecological fieldwork in a white water, inundation forest on the Solimoes River
near Manaus (Adis 1977 and 1979). Here the season can be divided into an
inundation phase, when the forest may flood to a depth of 4 m, and a non-inun-
dation phase. The inundation phase occurs March to August when the receding
waters mark the end of the inundation phase and the beginning of the non-inun-
dation phase. The latter begins with a dry period that lasts until December when
the rainy period of the non-inundation phase begins the accumulation of waters
for the beginning new inundation phase in March.

Fruits of the tree Pseudobombax munguba were found to be the main source
of nutrition for D. urbahni during the dry season of the non-inundation phase.
These trees lose their leaves during the beginning of the inundation phase, and
by June and July produce flowers and leaf buds, the fruit being formed and
dropped from the trees between August and October—the end of the inundation
phase until the beginning of the non-inundation phase.

Adults of D. urbahni appeared on the forest floor during the first week of the
non-inundation phase (the dry season); there they mated, usually near fallen
‘‘Munguba’’ fruits. The females laid their eggs in chapped fruits or on the soil
beneath and beside open or still closed fruits. Apparently fruits in the canopy

VOLUME 95, NUMBER 2 375

were not attacked. After 5-6 days first instar nymphs could be found; new gen-
eration adults began to appear after about 6 weeks [compare with the 42-55 days
reported for D. delauneyi Lethierry by Ballou 1906]. One chapped fruit may
contain as many as 1000 nymphs of all instars, while the average was 400 nymphs
per fruit (=61)!

Dead adults of the former generation were found near *‘Munguba’’ fruits 3-4
weeks after the forest dried up. At this time, the gregarious nymphs, migrating
on the forest floor, found newly dropped and already chapped fruits and fed on
the contained seeds. Field and laboratory experiments found that newly hatched
nymphs and adults preferred ripe but still soft seeds over dry or hard seeds; and
if no seeds were available they would feed on dead arthropods. Eight to ten
weeks after the forest had dried up, nearly all nymphs had transformed to adults
which frequented herbs, shrubs and young trees of the forest for another 2-3
weeks but then disappeared until the end of the dry season in November. Within
the rainy season of the non-inundation phase (December until March), adults
were seen only sporadically.

For the period from December to August, which also includes the inundation
phase, the source of nutrition for these insects is unknown. Pseudobombax mun-
guba is restricted to white water, inundation-forest (“‘Varzea’’); hence at other
localities, e.g., along blackwater rivers or on “‘terra firme’’ areas, different plants

or “‘Munguba’”’ species must provide reproductive sites and food sources for D.
~ urbahni.

Previous publications reported Bombacaceae to be visited by at least nine other
Dysdercus species (comp. Doesburg 1968, pp. 184-189; Silva et al. 1968 pp. 53-
56): D. fernaldi fuscofasciatus Blote (Bombax globosum; Suriname), D. fulvon-
iger (De Geer) (Bombax globosum; Suriname), D. fulvoniger discolor Walker
(Bombax malabaricum), D. honestus Blote (‘“‘Sumauma,”’ Brazil), D. longirostris
Stal (‘Sumatma,”’ Brazil), D. maurus distant (Bombax globosum; Suriname),
D. mimus (Say) (Bombax globosum (‘‘Sumatma’’); Belém, Brazil). D. mimus
distant Blote (sucking on seeds of ‘‘Sumauma’’ which had fallen to the floor;
Para, Brazil), D. ruficollis (Bombax ventricosa) and Dysdercus spp. (Bombax
discolor).

Acknowledgments

We are grateful to W. L. Overal (MPEG, Belém) and Norman Penny (INPA,
Manaus) for providing us with specimens. The manuscript was reviewed by Nor-
man Penny and typed by Irmgard Adis, who joined the field excursions, and
Silver B. West (Smithsonian Institution, Washington). We heartily thank Amy
Levin (S.I., Washington) for the literature research by computer.

Literature Cited

Adis, J. 1977. Programma minimo para analises de ecossistemas: Arthropodos terrestres em florestas

inundaveis da Amazonia Central.—Acta Amazonica 7(2):223—229.

. 1979. Vergleichende Okologische Studien an der terrestrischen Arthropodenfauna zentral

amazonischer Uberschwemmungswalder.—Ph.D. thesis, Univ. Ulm (West Germany), 99 pp.

Ballou, H. A. 1906. Cotton Stainers.—West Indian Bull. 7:64—-85.

Beccari, F., and V. Gerini. 1970. Catalogo delle Specie appartenenti al genere Dysdercus Boisduval
(Rhynchota, Pyrrhocoridae).—Revista di Agricolture subtropicale e tropicale 64:20-72.

376 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Doesburg Jr., P. H. van. 1968. A revision of the New World species of Dysdercus Guérin Méneville
(Heteroptera, Pyrrhocoridae).—Zoologische Verhandelingen 97:1—215.

Lima, A. M. da Costa, N. Guitton, and O. V. Ferreira. 1962. Sobre as espécies americanas do género
Dysdercus Boisuval (Hemiptera, Pyrrhocoridae, Pyrrhocorinae).—Memorias do Instituto Os-
waldo Cruz 60(1):21-57.

Silva, A. G. d’Aratjo, C. R. Congalves, D. M. Galyao, A. J. L. Goncalves, J. Gomes, M. do
Nascimento Silva, and L. de Simoni. 1968. Quarto catalogo dos insetos que vivem nas plantas
do Brasil, seus parasitos e predadores.—II(1):622 pp., Rio de Janeiro.

(JA) Max-Planck-Institut fur Limnologie, AG Tropenokologie D-232 Plon (Hol-
stein), West Germany in cooperation with Instituto Nacional de Pesquisas da
Amazonia (INPA), Manaus, Amazonas, Brazil; (RCF) U.S. National Museum of
Natural History, Smithsonian Institution, Washington, D.C. Present address:
INPA, c.p. 478, 69000 Manaus, Amazonas, Brazil.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 377-385

TWO NEW SPECIES OF THE FROG GENUS HYLODES
FROM CAPARAO, MINAS GERAIS, BRASIL
(AMPHIBIA: LEPTODACTYLIDAE)

W. Ronald Heyer

Abstract.—Two new species of the leptodactylid frog genus Hylodes, H. babax
and H. vanzolinii, are described from the Parque Nacional do Caparao, Minas
Gerais, Brasil. The identities of Elosia (=Hylodes) glabra and H. lateristrigatus
are reviewed.

During a survey of the herpetofauna of the Parque Nacional do Capara6o in
November—December 1980, Dr. P. E. Vanzolini and I collected specimens of two
species of Hylodes. Study of these frogs in the laboratory reveals that both
Species are new to science.

Examination of other Hylodes collected recently in the Atlantic Forests of
Brasil, suggests that the current allocation of specific names to certain Hylodes
Species is in error. Reassignment of specific names is clarified preparatory to the
_ description of the two new species from Caparao.

Advertising calls were recorded on a Uher CR 134 cassette recorder with a
Uher microphone or a Sony TCM 280 cassette recorder with a Sennheiser K3U
microphone. Calls were analyzed on a Kay Sonagraph 6061 B, narrow filter, AGC
in the off position.

Current Status of Names

Relatively few names have been proposed for members of the genus Hylodes.
As currently understood, the following clusters of species can be recognized on
the basis of external morphology. (1) The AH. pulcher group contains Hylodes
pulcher (B. Lutz, 1951), a very distinctive, moderate sized, slender, ranoid-like
species. The distinctiveness of the species derives from its bright life colors; most
individuals have bright blue and/or yellow dorsal spotting. (2) The H. mertensi
group contains Hylodes mertensi (Bokermann, 1956), a large (56 mm SVL), ro-
bust species with leathery dorsal skin. (3) The H. nasus group contains species
that are moderate to large sized, of robust body form, with granular dorsal sur-
faces, and lacking light dorsolateral stripes. Eight names have been proposed for
members of this group: Elosia aspera Miller, 1924; Hyla nasus Lichtenstein,
1823; Hyla ranoides Spix, 1824; Elosia nasuta Tschudi, 1838; Elosia bufonium
Girard, 1853; Hylodes truncatus Steindachner, 1864; Elosia nasus meridionalis
Mertens, 1927; and Elosia perplicata Miranda-Ribeiro, 1926. (4) The H. lateri-
strigatus group contains species that are of small to moderate size, body form
slender, ranoid-like, dorsum smooth, and in most members with light dorsolateral
stripes. The names proposed for this group are Elosia lateristrigata Baumann,
1912; Elosia glabra Miranda-Ribeiro, 1926; Elosia magalhaesi Bokermann, 1964;
Elosia ornata Bokermann, 1967; Hylodes regius Gouvéa, 1979. Both new species
belong to this latter cluster.

378 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

4 FOF ORE GF! POLES AF! PEGS;

| (22
TIME IN SECONDS

Fig. 1. Advertising call of Hylodes lateristrigatus. From specimen USNM 208557, Brasil, Rio de
Janeiro, near Teresopolis. Recorded 8 December 1977, air temperature 21.2°C.

The remainder of this section discusses the status of the names in the H.
lateristrigatus group and summarizes the advertising calls of those species for
which calls are known to facilitate comparison with the two new species.

Bokermann (1967b) discussed the taxonomic status of Elosia (=Hylodes) lat-
eristrigata and glabra. Prior to Bokermann’s 1967 papers, only one species was
recognized in the H. lateristrigatus group, i.e., the oldest available name, /ater-
istrigatus. Bokermann (1967b) discerned two distinctive advertising calls asso-
ciated with specimens from Friburgo in the Organ Mountains and Paranapiacaba,
Sao Paulo, respectively. He allocated the name Jateristrigatus to the Organ
Mountain species.

Recent collections from around Teresopolis (also in the Organ Mountains) doc-
ument the presence of two syntopic members of the H. lateristrigatus group. The
two species are similar morphologically and differ most markedly in size. The
larger species (males 38-39 mm SVL) has a call of about 1.5 s duration which
consists of about 13 notes; each note is a rising whistle with a dominant frequency
of 3700-4300 Hz with about 9 notes per second (Fig. 1). The smaller species
(males 33-34 mm SVL) has a call of about 1.1—1.3 s duration which usually
consists of only 2-4 notes; each note is a rising and falling whistle with a dominant
frequency of 4300-5500 Hz, with 3-4 notes per second (Fig. 2). Bokermann
(1967b) assigned the name E. Jateristrigata Baumann, 1912, to the species with
the call shown in Fig. 2 (compare with Fig. 1 in his paper; differences are due to
means of analysis, his figures emphasizing harmonic structure).

The type-locality given by Baumann is simply “*‘Orgel-Gebirge.’’ Baumann’s
(1912) description contains no measurements, although his Figure 1 on Plate 4 is
drawn at natural size. I measured the illustration as 38.9 mm SVL, which agrees
with data for the larger species at Teresopolis. This form always has sharply
defined dorsolateral and lower lip-upper arm stripes, as shown in Baumann’s
figure. Thus, H. lateristrigatus (Baumann) refers to the larger species, with the
call shown in Fig. 1, contra Bokermann (1967b).

The species that Bokermann (1967b) recorded from Paranapiacaba and referred
to as glabra has a call duration of about 2 seconds with 15-30 notes per call, each
note a rising whistle with a dominant frequency of 5600-6000 Hz (although his
Fig. 2 shows a dominant frequency of about 4700-5500 Hz), and with 10-13 notes

VOLUME 95, NUMBER 2 sg)

| 2

TIME IN SECONDS

Fig. 2. Advertising call of Hylodes sp. From specimen USNM field 6124, Brasil, Rio de Janeiro,
near Teresopolis, calling from same stream as USNM 208557. Recorded 8 December 1977, air tem-
peratune 21. 22C-

per second. Bokermann examined the type of Elosia glabra Miranda-Ribeiro, but
was not able to locate the specific site of “‘ribeirao da Passagem’’ within the
general type locality of Itatiaia. However, Bokermann found a species of the H.
lateristrigatus group on the lower slopes of Itatiaia at 700 m which compared well
_ with the specimens from Paranapiacaba and the type of E. glabra. Two facts
suggest that E. glabra does not refer to the species that Bokermann recorded
from Paranapiacaba and collected from the lower slopes of Itatiaia. The ‘‘ribeirao
da Passagem’”’ is in the planalto of Itatiaia at 2200 m elevation at approximately
22°25'S, 44°39’W (P. E. Vanzolini, in litt.). Second, the figures of E. glabra
(Miranda-Ribeiro 1926, Pl. 4, Figs. 1, la, 1b) do not appear to represent a species
of the H. lateristrigatus group. The figures in Miranda-Ribeiro’s work are accu-
rate representations of the specimens and species they represent, and show E.
glabra as a completely uniform brown frog. There are two species of frogs in the
upper elevations of Itatiaia that have occasional specimens with a uniform pat-
tern: Hylodes pulcher and Eleutherodactylus (Basanitia) nigriventris. The disks
of the figure of E. glabra appear Basanitia-like; the snout shape is Hylodes-like.
The possibility that E. glabra refers to an upper elevation frog, rather than a
lower elevation Hylodes is merely pointed out here and resolution of this problem
is deferred until it can be dealt with in a revision of the entire genus.

The conclusions drawn from the above comments are: (1) Hylodes lateristri-
gatus 1s the Organ Mountain species with a fast call rate with more notes; (2) the
species Bokermann referred to as Elosia lateristrigata currently has no name;
(3) the species name glabra may not refer to the species for which Bokermann
described and figured the call; and (4) the calls of H. lateristrigatus (in the sense
used here) and the calls described by Bokermann (1967b) as glabra are distinctive
and most probably represent two distinct species.

Hylodes magalhaesi is a moderate sized species (male 30 mm SVL), some
individuals of which have a distinctive belly pattern of discrete light spots on a
dark background. The advertising call averages about | s duration, with about 26
notes per call; each note has a dominant frequency centering on about 2000 Hz,
and a rate of 20-25 notes per second (Bokermann 1964).

Hylodes ornatus is a small species (males 25 mm SVL) (Bokermann 1967a).

380 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

i a ee 2 0 ee OO 2 Oe oO 2 a

TIME IN SECONDS

Fig. 3. Advertising call of Hylodes regius. From specimen USNM field 5316, Brasil, Rio de
Janeiro, Brejo da Lapa, Itatiaia. Recorded 7 January 1977, air temperature 18—20°C.

Most individuals have a distinctive dorsal pattern of two broad light dorsal stripes
in addition to the narrow light dorsolateral stripes. Males have external vocal
sacs indicating the species is vocal, but the call is presently unrecorded.

Hylodes regius is a moderate sized species (males 33-36 mm SVL) with a
distinctive dorsal pattern of small, irregular brilliant yellow spots on the dorsum
(Gouvéa 1979). The call is reported here for the first time. The call duration is
about 1.7 s, with about 22-32 notes per call. Each note is a sharply rising whistle
with a dominant frequency ranging from 5200-6300 Hz, and with 14-19 notes per
second (Fig. 3).

The two new species from Caparao are distinct from all taxa discussed above.

Hylodes babax, new species
Fig. 4

Holotype.—MZUSP 57949, adult male, from Brasil: Minas Gerais; Parque Na-
cional do Caparao, 1200 m, 20°26’S, 41°47’W. Collected by W. Ronald Heyer, 30
November 1980. |

Paratopotype.—USNM 222553, male, same data as holotype.

Diagnosis.—The dorsal coloration is sharply set off from the lateral coloration
by a weakly developed light dorsolateral stripe. This pattern distinguishes Hy-
lodes babax from the other new species from Caparao. Hylodes babax is small
(male SVL 31-33 mm) and has a dark belly with light spots; the two species at
Teresopolis are larger (/ateristrigatus male SVL 38-40 mm; the second species
male SVL 33-34 mm) and have light bellies with dark spots. Hylodes babax is
larger than Hylodes ornatus (males SVL 25 mm) and lacks the distinctive dorsal
pattern of a pair of elongate light dorsal bands characteristic of H. ornatus.
Hylodes babax resembles H. magalhaesi from Campos do Jordao, Serra da Man-
tiqueira, Sao Paulo, Brasil, in size and general coloration. In life, the under
surfaces of the legs are brick red in H. babax, yellowish in H. magalhaesi. The
advertising calls of H. babax and magalhaesi are strikingly different (compare
Fig. 4 of Bokermann 1964, with Fig. 5 here) .

Description of holotype.—Snout rounded-truncate from above, protruding in
profile; canthus rostralis angular-obtuse; loreal weakly concave in cross section;

VOLUME 95, NUMBER 2 381

Fig. 4. Holotype of Hylodes babax (MZUSP 57949, a male): dorsal and ventral views.

tympanum distinct, diameter about 4 eye diameter; vomerine teeth in two small
transverse patches, medial and just posterior to choanae; vocal slits and well
developed lateral vocal pouches present; first finger long, just longer than second;
thumb lacking nuptial asperities; dorsal texture essentially smooth, with small
pebble like granulations; weak supratympanic fold, no other body folds obvious;
no body glands; belly smooth, ventral femur granular; fingers and toes disked,
disks about 1% again as broad as toe width just behind disks, toe disks slightly
larger than finger disks, each disk with a pair of dorsal scutes; fingers and toes
with considerable fringe; subarticular tubercles present but not prominent; ovate
inner metatarsal tubercle not quite twice as large as semicircular outer metatarsal
tubercle; well developed tarsal fold extending about % distance of tarsus, con-
tinuous distally with outer toe fringe of toe 5; no metatarsal fold; outer tarsus and
sole of foot smooth.

SVL 30.6; head length 11.6; head width 9.7; eye—nostril distance 2.5; femur
15.1; tibia 17.2; foot 15.9 mm.

Dorsal pattern of variegated brown and tan markings with irregular dark brown
interocular mark; sides almost black, demarcated from dorsal coloration by thin
light pinstripe extending from tip of snout over the eye to the upper groin; light
lip stripe, broader but less distinctive anteriorly; upper limbs brown with irregular
dark transverse bars, upper forearm with light pin stripe on anterior aspect of
arm from shoulder to inner elbow; throat black with very few, somewhat sym-
metrically arranged light dots; belly black with contrasting white irregular spots,
posterior surface of thigh with indistinct dark mottle; ventral surface of thigh with
pale straw colored patch, other ventral limb surfaces black with little pale areas.

In life the upper lip stripe tannish bronze; the belly with contrasting black and
white markings, and the lower surface of the thigh brick (not brilliant) red.

382 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

{feeebbe

| 2
TIME IN SECONDS

Fig. 5. Advertising call of Hylodes babax. From specimen USNM 222553, recorded 30 November
1980, 1540 h, air temperature 21.6°C.

Description of paratopotype.—The paratopotype resembles the holotype in
most aspects with the following exceptions: dorsal coloration not as dark, weak
bronze colored dorsolateral fold demarcates dorsal and lateral patterns; belly with
small and less distinct light spots; SVL 32.6 mm.

Aavertising call.—Call duration 0.23—0.48 s, mean 0.36 s; 4-8 notes per call,
mode 6; each note a rising whistle, typically first note lower in pitch (mean
dominant frequencies 4290-5030 Hz) than remaining notes (mean dominant fre-
quencies 4690-5420 Hz); notes produced at a mean rate of 16.2 notes per second
(Fig. 5).

Etymology.—TYhe name babax is Greek for chatterer, in allusion both to the
insistent diurnal calling of this species (as in many other Hylodes species) and to
the fact that of the two species of the H. lateristrigatus group from Capara6o, this
is the only one known to call.

Hylodes vanzolinii, new species
Fig. 6

Holotype. —MZUSP 57950, male, from Brasil: Minas Gerais; Parque Nacional
do Caparao, 2300 m, 20°26’S, 41°47’W. Collected by W. Ronald Heyer and P. E.
Vanzolini, 5 December 1980.

Paratopotypes.—MZUSP 52923, collected by Lynn C. Branch, November
1977; USNM 222554-5, same data as holotype.

Diagnosis.—TYhe dorsal color pattern is not sharply set off from the lateral
coloration. This pattern is distinct from all other members of the H. lateristrigatus
group with the possible exception of glabra (see comments in previous section),
which in the original figure is completely uniform dorsally and ventrally. Hylodes
vanzolinii has small bright yellow dorsal spots in life (white in preservative) and
a variegated belly, differing from the figures of E. glabra.

Description of holotype.—Snout rounded-truncate from above, protruding in
profile; canthus rostralis angularly obtuse; loreal slightly concave in cross section;
tympanum about /% diameter of eye; vomerine teeth in two small slightly trans-
verse patches between posterior extent of choanae; no vocal slits or vocal sac;
first finger long, just longer than second; thumb lacking nuptial asperities; dorsal

VOLUME 95, NUMBER 2 383

Fig. 6. Holotype of Hylodes vanzolinii (MZUSP 57950, a male): dorsal and ventral views.

texture smooth; very weak supratympanic fold, otherwise no other body folds or
glandular structures; belly smooth, under surface of thighs granular; finger and
toe tips with disks, disks about % again as broad as digit immediately behind
disk, finger and toe disks about equal size, upper surface of disks with a pair of
scutes; fingers with lateral ridge, toes with extensive lateral fringe; subarticular
tubercles moderate; inner ovate metatarsal tubercle much larger than rounded
outer metatarsal; extensive tarsal fold extending about 7% distance of tarsus, con-
tinuous distally with toe fringe on outer side of first toe; no metatarsal fold; outer
tarsus and sole of foot smooth.

SVL 29.0; head length 10.4; head width 9.5; eye—nostril distance 2.0; femur
16.0; tibia 16.9; foot 15.8 mm.

Dorsum almost black with nondescript grayish mottling and with a few small
light dots; dorsal pattern not distinct from lateral pattern except on head with an
irregular whitish gray stripe from tip of snout to just over tympanum demarcating
dorsal pattern from uniform dark lateral head pattern; light upper lip stripe in-
complete, weaker in front of eye than behind; dorsal limb surfaces almost uni-
formly dark brown, front of forearm with very weak, incomplete light pin-stripe;
throat mostly brown with a symmetrical series of medial coalescing spots; belly
variegated brown and white in about equal ratio; posterior and ventral surfaces
of thighs uniform brown.

In life upper lip stripe dull white; dorsum dark green-black with yellowish
flecks; belly white and tan; posterior surface of thigh uniform tan.

Description of paratopotypes.—USNM 222554, a 37.3 mm female, resembles
the holotype in pattern with the following differences: the upper lip has a series
of light dots, not a stripe; there is no light stripe separating the dorsal color from
the side color on the head; the throat is mostly white with some dark markings.
MZUSP 52923 is uniformly dark dorsally and laterally except for a few small light
dorsal spots and a faint light upper lip stripe mostly in front of the eye; the belly
is dark with small, distinct light spots.

384 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Advertising call.—The fact that no Hylodes vanzolinii were heard calling, to-
gether with the lack of vocal slits in the males, strongly suggests that this species
lacks an advertisement call.

Etymology.—It is a pleasure to name this species for P. E. Vanzolini. First, I
wish to commemorate an enjoyable field trip during which he shared many in-
sights concerning the Atlantic Forest with me. Second, I wish to pay tribute to
Dr. Vanzolini’s studies on the montane lizard fauna of eastern Brasil where he
developed the concept of stranded species. Without his studies, explanation of
the history of this new montane frog would not be possible.

Discussion

Aylodes vanzolinii replaces H. babax altitudinally in the Parque Nacional do
Caparao, with the latter occurring along streams in the Atlantic Forest vegetation
and Hf. vanzolinii occurring along the exposed streams at elevations above the
Atlantic Forest vegetation (1900 m and above). The distributions of the species
are so far known only from the collections reported here.

Hylodes babax is similar to several other members of the H. lateristrigatus
group and on the basis of color pattern is predicted to be related to one of these
species than to the altitudinally adjacent H. vanzolinii.

Hylodes vanzolinii and H. regius (Itatiaia, Serra da Mantiqueira) both have
small yellow spots on the dorsum. Hylodes vanzolinii differs from H. regius in
the dorsolateral stripes and in lacking a voice. These two species do not seem to
be closely related.

As far as is known, there are only two Hylodes species that lack advertising
calls, H. pulcher and vanzolinii. Both of these species are montane (in the coastal
mountain system of Brasil at elevations above 1700-1900 meters) species that
have striking dorsal colors in life. The relationships between H. pulcher and
vanzolinii are probably not close, however, as the color patterns and sizes are
very distinctive.

There are two major zoogeographic patterns that account for montane distri-
butions in eastern Brasil. The first is one of relictual distribution of stranded
Species associated with drier and cooler climates that today are restricted to the
upper elevations of mountains, but that were much more widespread during gla-
cial maxima periods of the Pleistocene (Vanzolini 1982; Vanzolini and Ramos
1977). The montane species that exhibit this zoogeographic pattern are closely
related to each other. The second pattern is one of in situ differentiation, where
the present montane species were independently derived from lowland forms.
Hylodes vanzolinii appears to fit into this second zoogeographic pattern. Detailed
resolution of relationships awaits biochemical and morphological analyses.

Acknowledgments

P. E. Vanzolini (Museu de Zoologia da Universidade de Sao Paulo, MZUSP)
obtained the proper Instituto Brasileiro de Desenvolvimento Florestal permits
and invited me to join in the exploration of the herpetofauna of the Parque Na-
cional do Caparao. George R. Zug and Ronald I. Crombie (National Museum of
Natural History, Smithsonian Institution, USNM) critically read the manuscript.
George Steyskal (USNM) advised on scientific name usage and formation.

VOLUME 95, NUMBER 2 385

The following programs and institutions supported this research: Fundacao de
Amparo a Pesquisa do Estado de Sao Paulo; Museu de Zoologia da Universidade
de Sao Paulo; and the International Environmental Science Neotropical Lowland
Research Program, Smithsonian Institution.

Literature Cited

Baumann, F. 1912. Brasilianische Batrachier des Berner Naturhistorischen Museums.—Zoologischen
Jahrbtichern, Abteilung fiir Systematik, Geographie und Biologie der Tiere 33:87-172 + plates
4&5.

Bokermann, W. C. A. 1964. Una nueva especie de Elosia de la Serra da Mantiqueira, Brasil (Am-

phibia, Leptodactylidae).—Neotropica 10:102-107.

. 1967a. Una nueva especie de Elosia de Itatiaia, Brasil (Amphibia, Leptodactylidae).—Neo-

tropica 13:135-137.

. 1967b. Notas sobre cantos nupciais de anfibios brasileiros. II: O canto de ‘‘Elosia lateristri-

gata’’ e ‘‘Elosia glabra’ (Anura).—Revista Brasileira de Biologia 27:229-231.

Gouvéa, E. 1979. Uma nova espécie de elosiineo da Serra do Itatiaia (Amphibia, Anura, Leptodac-
tylidae).—Revista Brasileira de Biologia 39:855-859.

Miranda-Ribeiro, A. 1926. Notas para servirem ao estudo das Gymnobatrachios (Anura) Brasileiros—
Tomo primeiro.—Archivos do Museu Nacional do Rio de Janeiro 27:1—227 + 22 plates.
Vanzolini, P. E. 1982. A new Gymnodactylus from Minas Gerais, Brasil, with remarks on the genus,
on the area and on montane endemisms in Brasil (Sauria, Gekkonidae).—Papéis Avulsos de

Zoologia 34:403—413.

,and A. M. M. Ramos. 1977. A new species of Colobodactylus, with notes on the distribution

of a group of stranded microtetid lizards (Sauria, Teiidae).—Papeéis Avulsos de Zoologia 31:19-

47.

Amphibians and Reptiles, Department of Vertebrate Zoology, Smithsonian In-
stitution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 386-391

A NEW CAVE PLATYMANTIS (AMPHIBIA: RANIDAE)
FROM THE PHILIPPINE ISLANDS

Walter C. Brown and Angel C. Alcala

Abstract.—A new cave-dwelling frog, Platymantis spelaeus (Ranidae), is de-
scribed from southern Negros Island in the Philippines. This is the second cave-
dweller and the twelfth species of the genus known from this archipelago.

Recognized species of the genus Platymantis include two from the Fiji Islands,
nine from the Solomons, nine from the Bismarcks, four or five from New Guinea
and small associated islands, one from the Palaus, and eleven from the Philip-
pines. With the exception of one species (as presently defined) which ranges from
New Guinea to the Solomons, all are endemic to the regions indicated.

In the western Philippines the genus is absent from Palawan and associated
small islands. In the rest of Philippines, three species (P. corrugatus, P. dorsalis
and P. guentheri) are widespread, being recorded from Mindanao in the south
through Luzon in the north. In addition, P. ingeri occurs in Mindanao and other
islands in the southern part, P. hazelae has been recorded from Negros and
Luzon Islands, and the very closely related P. polillensis is known from Polillo
Island. The remaining five species are known from limited localities on single
islands. Platymantis cornutus and P. subterrestris are recorded from Luzon Is-
land, P. lawtoni and P. levigatus from Tablas Island to the south of Luzon and
east of Mindoro, and P. insulatus from the very small island, Gigante South, off
the northeast coast of Panay.

Typically these frogs inhabit the rain forest, being either arboreal or in habitats
of the duff or under rocks and logs on the forest floor. Only one species, P.
insulatus (Brown and Alcala, 1970), has previously been recorded as cave-dwell-
ing. Now a second such species is described in this paper. The type series is from
two limestone caves in a forested area in southern Negros Island. 3

Measurements of specimens were made using a Helios dial caliper. Head length
is measured from the tip of the snout to the posterior edge of the tympanum and
head breadth at the widest point near the angle of the jaws. The diameter of the
eye is from the anterior to the posterior edge of the socket. Finger and toe lengths
are to the proximal edge of the proximal subarticular tubercle. Other measure-
ments are probably not subject to much variation in method.

Platymantis spelaeus, new species
Fig. 1

Holotype.—California Academy of Sciences number 153469, mature female,
collected in a limestone cave, Tiyabanan Barrio, Basay, southern Negros Ori-
ental, Negros Island, Philippines, by Angel Alcala and C. A. Ross, 18 March
1981.

Paratypes.—California Academy of Sciences 153470-83; United States Na-
tional Museum 221838-39; Australian Museum, Sydney R98394; Field Museum

VOLUME 95, NUMBER 2 387

Fig. 1. Platymantis spelaeus, showing the lack of ridges on the dorsum, the large tympanum, and
characteristic features of the color pattern.

Natural History 213331; British Museum (Natural History) 1981.9; Museum of
Comparative Zoology, Harvard A100300; Silliman University, Philippines 2288-
92; American Museum Natural History 109454, all from two caves in southern
Negros Island.

Diagnosis.—This Platymantis is distinguished from other Philippine species by
the following combination of characters: (1) Large size (snout—vent length 41.5-—
60.5 mm for 17 adults); (2) The only slightly dilated finger disks, about the same
diameter as those of the toes; (3) Tympanum relatively large, its diameter about
62-82% of that of the eye; (4) Skin of the dorsum rough (shagreened), neither
smooth nor with prominent tubercles or elongate ridges.

Description.—A large Platymantis, snout—vent length (SVL) 41.5-46.9 mm for
5 males and 52.8-60.5 mm for 11 females (Table 1); head breadth 107-111% of
head length; snout broadly rounded, its length 40-44% of head breadth and 41-
49% of head length; upper jaw protruding; canthus rostralis rounded; lores mod-
erately oblique, concave; diameter of eye 27-37% of head breadth and 63-88%
of snout length; tympanum large, its diameter 63-82% of eye diameter, 105—147%
of interorbital distance, and about twice the diameter of the third finger disk;
prominent fold dorsal and posterior to tympanum; dorsum shagreened (Fig. 1)
but lacking ridges; venter smooth; fingers slender, without webs; tips of fingers
with rounded disks having a circummarginal groove but lacking a transverse
ventral groove (Fig. 2); diameter of disk of third finger 20-25% of length of third
finger (measured to base of second subarticular tubercle); disk of second finger

388 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Snout-—vent lengths for Philippine species of Platymantis (number in sample in paren-

theses).

al

~~ Males

Snout—vent length (mm)

Sex

Species undetermined Females
P. cornutus 30.7 (1)
P. corrugatus 28.5-35.0 (15) 33.6—-50.2 (15)
P. dorsalis 21.5—30.5 (25) 25.5—42.4 (30)
P. guentheri 27.4-35.5 (24) 38.3-49.1 (20)
P. hazelae 20.6—27.6 (15) 20°7—31-7 G5)
P. ingeri 23.0-30.5 (16) 26.9-34.5 (23)
P. insulatus 37.8-41.7 (3) 40.2-45.5 (3)
P. lawtoni — 39.0 (1)
P. levigatus 30.3 (1) 38.5 (1)
P. polillensis 20.2—22.6 (2) a
P. spelaeus 41.5-46.9 (6) 52.8-60.5 (11)
P. subterrestris 24.0-26.3 (2) 21 Dl)

nearly as large as that of third finger (Table 2); first finger slightly longer than
second when adpressed; fingers with prominent rounded subarticular tubercles;
a prominent palmer tubercle at base of each finger; three large metacarpal tuber-
cles; hind limbs relatively long, length of tibia 52-63% of snout—vent length and
127-148% of head breadth; toes with small web at base; tips dilated into rounded
disks with circummarginal grooves; diameter of disk of third toe 87—1.07% of
diameter of disk of third finger; subarticular tubercles prominent, round-pointed;

Table 2.—Proportions in samples of some of the larger Philippine species of Platymantis.

P. guentheri P. insulatus P. levigatus P. spelaeus
(n = 20) (n = 5) (n = 2) (n = 19)

Eye diameter
EL 0.33-0.39 0.37-0.41 0.29-0.30 0:97-0:37
Eye. diameter, 1.04-1.09 0.99--1.06 0.97-0.98 0.63-0.88
Snout length
Snout length
Tee 0.30-0.35 0.35-0.39 0.29-0.31 0.40-0.49
os
Tympanum diameter 0.37-0.48 0.48-0.61 0.33-0.40 0.63-0.82
Eye diameter
Tympanum diameter
PRM DMEN CUTE gi 0.63-0.87 122s 1.40-1.50 1.89-2.60
:
dutevorbits bread 0.45-0.67 0.44-0.49 0.60-0.61 0.47-0.67
Eye diameter
3 .
Bred eulicor sdipenigisk 0.36-0.51 0.27-0.31 0.17-0.20 0.20-0.25
Length of 3rd finger
Breadth of 2nd finger disk
See rE Re 0.73-0.90 0.63-0.74 0.91-1.00 0.76-0.95
‘i

Readih oPSnC motes 0.47-0.56 0.62-0.74 0.85-0.90 0.87-1.07

Breadth of 3rd finger disk

VOLUME 95, NUMBER 2 389

Fig. 2. P. spelaeus (SU 2288): palmar view of hand.

an elongate outer metatarsal tubercle and a rounded inner one; solar tubercles
lacking (Fig. 3).

Measurements (in mm) of holotype.—Snout-vent length 60.5; head length 22.5;
head breadth 24.4; snout—length 10.0; eye diameter 8.8; tympanum diameter 5.5;
interorbital distance 4.4; 3rd finger length 10.8; diameter of 3rd finger disk 2.7;
diameter of 3rd toe disk 2.5; hind limb length 95+; tibia length 31.5.

Color.—In preservative the dorsum and the head are mottled grayish-olive and
black with a short, broad, blackish *‘W”’ in the forelimb region. The lateral sur-
faces are more grayish with the darker mottling reduced. The lips are dark barred.
The dark transverse bands on the limbs are about as wide as or wider than the
light bands (Fig. 1). The venter is creamy white with some faint gray mottling.

In life the dorsum is olive-green to brown with darker mottling. The upper
surfaces of the thighs are dark barred and the inner surface either orange or
lavender. The venter is cream colored with or without brown flecks.

Comparisons.—This is the largest species of the genus known from the Phil-
ippines (Table 1). Eight of the previously described species are small, arboreal
forms with the SVL in mature males under 36 mm and with females rarely at-
taining a SVL greater than 40 mm (P. dorsalis does). Only one mature female is
known for both P. lawtoni and P. levigatus, in which the SVL is 39 and 38.5,
respectively, and it is possible that a SVL of 40 mm may be exceeded in those
species. Only males of P. insulatus overlap P. spelaeus in SVL and only females

390 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. P. spelaeus (SU 2288): plantar view of foot.

of P. corrugatus, P. dorsalis, P. guentheri and P. insulatus approach females of
P. spelaeus in size although they do not overlap (Table 1).

Most Philippine species also have broadly dilated digital disks, especially on
the fore limbs (Inger 1954; Brown and Alcala 1963, 1970, 1974). Only in P. cor-

VOLUME 95, NUMBER 2 391

rugatus, P. dorsalis and P. levigatus are the finger tips only slightly dilated as is
characteristic of P. spelaeus.

The shagreened skin of the dorsum (not smooth or with prominent tubercles
or ridges) is also characteristic of P. insulatus and P. lawtoni.

Platymantis spelaeus, insulatus and dorsalis are the most apt to be confused
in terms of general appearance. The somewhat smaller P. dorsalis has prominent,
short ridges on the dorsum, and the general color is more brown to reddish.
Platymantis insulatus is closer to P. spelaeus in size, texture of the skin on the
dorsum, and general color pattern, but the tips of the fingers are much more
broadly dilated and the tympanum is smaller (Table 2). The species P. spelaeus
and P. insulatus are probably closely related.

Ecological notes.—All specimens were collected near the entrances of two
moist limestone caves, in crevices or pockets in the walls or on the floor. Sixty
large (2.5—3.0 mm diameter) unpigmented eggs were found in the ovaries of a
large gravid female (55 mm SVL).

Acknowledgments

We gratefully acknowledge the assistance of Charles A. Ross of the U.S. Na-
tional Museum of Natural History, Smithsonian Institution in the field work and
W. Ronald Heyer of the same institution for the loan of specimens. We also thank
-Richard G. Zweifel, American Museum of Natural History, for reading the manu-
script.

Literature Cited

Brown, Walter C., and Angel C. Alcala. 1963. A new frog of the genus Cornufer (Ranidae) with

notes on other amphibians known from Bohol Island, Philippines.—Copeia 1963:672-675.

, and . 1970. A new species of the genus Platymantis (Ranidae) with a list of amphibians

known from South Gigante Island, Philippines.—Occasional Papers of the California Academy

of Sciences 84:1-8.

, and . 1974. New frogs of the genus Platymantis (Ranidae) from the Philippines.—

Occasional Papers of the California Academy of Sciences 113:1-12.

Inger, Robert F. 1954. Systematics and zoogeography of the Philippine Amphibia.—Fieldiana: Zo-
ology 33:183-531.

(WCB) Department of Herpetology, California Academy of Sciences, Golden
Gate Park, San Francisco, California 94118 and Menlo College, Menlo Park,
California 94025; (ACA) Department of Biology, Silliman University, Manila,
Philippines.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 392-397

A NEW SPECIES OF SPHAERODACTYLUS
(REPTILIA: SAURIA: GEKKONIDAE)
FROM EASTERN CUBA

Orlando H. Garrido and Albert Schwartz

Abstract.—A new species of gecko, Sphaerodactylus celicara, of the notatus-
group, is described from extreme northeastern Cuba. Comparisons are made with
its most closely related geographical congeners, S. notatus and S. bromeliarum.

A striking feature of the distribution of Sphaercdactylus in Cuba is that the
easternmost province, Oriente, harbors the greatest diversity of species, repre-
senting several species-groups of these small lizards. One of these, S. notatus,
belongs to a complex of geckos whose center of diversity is Hispaniola (and
secondarily Puerto Rico). On Cuba, there is only one species (S. bromeliarum
Peters and Schwartz) that is closely related to S. notatus, in contrast to 14 no-
tatus-group species on Hispaniola.

There are some peculiarities in scutellation and pattern in S. notatus on Cuba.
For instance, Keeling on the throat does not occur in specimens from the Isla de
la Juventud (=Isla de Pinos) or, on the main island, from Pinar del Rio to Ca-
magtey provinces. Throat keeling occurs more or less variably (in some individ-
uals from the same locality and in varying degrees) in Oriente specimens. A
second variable feature is the occurrence in females of a dark scapular patch with
2, rarely 1, included pale ocelli. In western and central Cuba, the female scapular
pattern is absent. In Oriente, on the other hand, 26% of the females have a
scapular patch and ocelli. Since absence of this patch, the presence of which is
characteristic of S. bromeliarum, is one of the distinctive characteristics of the
Cuban subspecies S. n. atactus Schwartz, these eastern specimens have seemed
anomalous. ?

The senior author sent a series of 14 specimens from the extreme northeastern
region of Cuba to the junior author. Many of these are larger lizards than typical
S.n. atactus and differ in scutellation; in addition, they to a large extent (but not
totally) clarify the problem of occurrence of a female scapular patch and ocelli.
The males normally have both the upperside of the head and throat vividly and
distinctly spotted or dotted with very dark brown. Data from these specimens
were combined with data from other geckos from this same general region, using
the presence of a scapular patch and ocelli and scutellar counts as criteria, and
we now have a clearer picture of the variation in notatus-like geckos in this
particular region. We have examined a total of 239 S. notatus from Cuba, in-
cluding 182 specimens from Oriente alone (on 118 of which full counts were
taken) and 37 from the Isla de la Juventud and the Archipiélago de los Canarreos,
and we feel secure in naming this northeastern population as a distinct species.

Sphaerodactylus celicara, new species

Holotype.—IZ 5613, adult female, from Asuncion, Maisi, Baracoa, Oriente
Province, Cuba, collected 10 May 1979 by Orlando H. Garrido.

VOLUME 95, NUMBER 2 393

Paratypes.—IZ 5611-12, IZ 5616-19, ASFS V44969-70, same data as holotype;
ASFS V44968, La Maquina, Gran Tierra, Maisi, Oriente Province, Cuba, 10 May
1979, M. L. Jaume; IZ 5378, Hotel Asuncion, Grand Tierra, Maisi, Oriente Prov-
ince, Cuba; 17 September 1978, P. Espinosa, H. Saragua, A. Vega; IZ 4725, 3
km W Yumuri, Oriente Province, Cuba, May 1976, Gilbert; IZ 5414-15, Zapote
de Mandinga, Baracoa, Oriente Province, Cuba, 10 May 1979, O. H. Garrido.

Associated specimens.—All are from Oriente Province, Cuba: IZ 3455-57,
Hoyos de Sabanilla; IZ 4214-16, La Poa, Sabanilla; IZ 3453, IZ 3537, IZ 3585,
base of Monte Iberia; IZ 3648, Jaguani, Baracoa: IZ 106, IZ 113, La Tinta, barrio
be Jauco, Baracoa; IZ 4096, La Yagruma, Maisi, Baracoa; IZ 4258, Punta de
Maisi; MCZ 13595-96, Jauco, seacoast at Cabo Maisi; MCZ 11215, Cueva de
Majana, Baracoa.

Definition.—A species of the notatus-group of Sphaerodactylus characterized
by the combination of: 1) moderate size, males to 32 mm, females to 31 mm
snout—vent length (SVL); 2) high number of dorsal scales (21-30) between axilla
and groin, each dorsal scale with 11-14 hair-bearing organs, each with one hair,
around the free edge and apex of the scale; 3) high number of scales (40-51)
around midbody; 4) scapular patch and ocelli almost always present in females,
only ocelli usually present in males; 5) heads of adults of both sexes heavily
dotted or marbled with dark brown, both dorsally, and ventrally on the throat,
in strong contrast to the brown color of the remainder of the body; 6) females

‘typically bilineate dorsally, males more or less unicolor to salt-and-pepper,
brown; 7) internasals 0—2, with a high incidence (35%) of internasals absent (=0).

Description of holotype.—Adult female, SVL 31 mm, tail length 34 mm. Scale
counts: dorsal scales keeled, tectiform, imbricate, between axilla and groin 29;
ventral scales between axilla and groin 24; scales around midbody 47; fourth toe
lamellae 11; 3/3 supralabials to mid-eye; no internasals; 2 postnasals. Throat,
chest, and ventral scales smooth, imbricate, cycloid. Dorsal color (preserved)
dark brown with dorsolateral pair of slightly paler lines (Fig. 1), strongly outlined
with black, beginning at well developed black scapular patch which includes two
pale (grayish) ocelli; head pattern basically dark trilineate, with a median dark
line, narrow on snout and expanding on occiput and continuing to scapular patch,
somewhat paler centrally, and on each side a narrow dark canthal line to the eye,
continuing posteriorly from eye as broad dark line to lateral margins of scapular
patch, the two forming a more or less square brownish area surrounding the
scapular patch. The lateral pair of cephalic lines bordered below by pale lines
that continue clearly above the forelimb insertion; pale area between lateral lines
and central dark line on head containing a dark postocular line extending to
occiput. Sides of neck brownish, dotted with darker brown; upper lip mottled
with strongly contrasting dark brown and pale ground color, the mottling ex-
tending onto the throat as strongly contrasting dark brown dots and lines,~this
pattern becoming diffuse on the chest which is stippled with dark brown. Ventral
scales edged with dark brown, resulting in a generally dark venter; tail brown
with obscure darker brown marbling; upperside of all limbs dark brown, stippled
or spotted with darker brown.

Variation.—The holotype and 30 paratypes and associated specimens show the
following variation. Largest males (IZ 5611, IZ 5617) 32 mm SVL (range 25-32
mm), largest females (IZ 5613, IZ 5619) 31 mm (22-31 mm); dorsals between
axilla and groin 21-30 (x = 25.8), ventrals between axilla and groin 22-34 (26.1);

394 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

R OVle

Fig. 1. Sphaerodactylus celicara, holotype (IZ 5613). Upper figure, dorsal view; middle figure,
lateral view of head pattern; lower figure, ventral view of throat pattern.

midbody scales 40-51 (45.2); fourth toe lamellae 9-13 (10.6; mode 11); suprala-
bials 3/3 (29 individuals), 3/4 (1), 4/4 (1); internasals 0 (11 individuals), 1 (17), 2
(3); male escutcheon 4—7 x 10-26; throat, chest, and ventral scales smooth in 23
specimens, throat scales keeled in 7 specimens; upper side of head lined or heavily
dotted in 7 males, head markings absent in 2; throats heavily dotted in 7 males,
these markings absent in 2; scapular patch and ocelli present in 21 females, absent
in one.

Of the four male paratypes, three agree well with the definition of the species.
They have the dorsum salt-and-pepper or apparently unicolor tan with strongly
contrasting dark brown head mottling or marking on a pale ground. The dark
dorsal head mottling continues onto the throat in these specimens as very heavy
dark mottling comparable to that of the upper surface of the head. Venters have

VOLUME 95, NUMBER 2 395)

20 22 24 26 28 30 32

SVL

Fig. 2. Histogram of snout—vent length (in mm) of two samples of S$. notatus (open rectangles)
and S. celicara (dark rectangles); males are shown above the horizontal line, females below it. Each
individual is indicated by an interval of 0.2 mm.

dark-edged scales. The exceptional male (IZ 5612, SVL 31 mm) lacks any dorsal
or ventral pattern. Although there is no dark scapular patch in males, two (IZ
5611, IZ 5617) have a faint pair of pale ocelli; the other two males (IZ 5612, ASFS
V44968) lack them.

Scapular patches and ocelli are present in all but one of the 10 female paratypes,
as well as in all 10 associated females and one juvenile. The dorsal bilineate
pattern is usually quite distinct (even in small lizards—IZ 5378, SVL 23 mm) and
is especially vivid in some (ASFS V44969). In some smaller lizards the tail has
a tiny black tip, with a broader white band proximally, which is in turn bounded
by a narrower black band. The head pattern is trilineate, with interspace dark
lines between the major head lines in all specimens; there is no blurring or dis-
tortion of the three major cephalic lines in large specimens. Scapular patches
vary from large and extensive (ASFS V44970) to relatively small and constricted
(IZ 5614), but they and the included ocelli are always present. Dark throat mark-
ings are almost always present and obvious, varying from heavy in the holotype
to less bold and lineate in some specimens. The distinct pale-and-dark neck
streaking is also a constant feature, although the more lateral nuchal lines may
disintegrate into a linear series of very dark spots or blotches (IZ 5618).

The associated specimens agree very well with our concept of S. celicara.
There are five males, ten females, and two juveniles (SVL 16 and 20 mm). Only
one male (IZ 3537) lacks a dotted or spotted head and throat, and another male
(MCZ 13595, SVL 25 mm) not only has a dotted head and throat but also has a
scapular patch and two ocelli, a most unusual condition. In the females, scapular
patches and ocelli are present in all but one (IZ 3453, adult, SVL 29 mm). The
juveniles show the female head and shoulder pattern.

396 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Extremes and means of four characters of S. notatus and S. celicara, plus data on
underbody keeling. S. notatus has been divided into two samples: Sample 1 from Pinar del Rio to
Camagiiey provinces, Sample 3 from Oriente Province.

S. notatus
Sample 1 Sample 2 S. celicara
n S71 118 30
Snout—vent length (mm) 6 20-30 20-30 25=3))
(25.8) (25.0) (28.7)
2 20-30 20-32 Mp =8))\
G55) (25.1) Q7T2)
Dorsal scales 18-28 18-27 21-30
CikG == 07) (22.1 + 0.4) (25.8 + 0.9)
Midbody scales 31-49 33-47 40-51
(37.4 + 1.4) (39.5 + 0.6) (45.2 = 1.2)
4th toe lamellae 7-11 i= 9-13
(9.3; M, = 10) (9.7; M, = 10) (10.8; M, = 11)
Ventral keeling
Throat none 2 (weak), 6 distinct 7 (absent in 23)
Chest none 1 (+ throat) none
Ventrals none 109 none

Comparisons.—Sphaerodactylus bromeliarum is similar in general appearance
but is not lineate dorsally, has keeled ventral scales, has a prominently black-
and-white banded tail tip in adults, and has smaller scales (midbody scales 56-58
versus 40-51 in S. celicara). The only known adult (female) of S. bromeliarum
has a SVL of 24 mm, much smaller than S. celicara.

It is with §. notatus that S. celicara most requires comparison. We have
grouped data on 57 specimens from Pinar del Rio to Camagtiey provinces (Sample
1) and 118 specimens from Oriente Province (Sample 2) for the purposes of dis-
cussion (Table 1, Fig. 2). Both samples of S. notatus average smaller than both
sexes of S. celicara in SVL. Samples 1 (21.6 + 0.7) and 2 (22.1 + 0.4) do not
differ significantly in dorsal scale counts, but they do differ significantly from S.
celicara (25.8 + 0.9). The means of midbody scale counts in Samples 1 and 2
(37.4 + 1.4 and 39.5 + 0.6) differ significantly from each other and they are also
significantly different from the mean in S. celicara (45.2 + 1.2). There are fewer
toe lamellae in the two samples of S. notatus than in S. celicara (Table 1).

Although the two samples of S. notatus as well as S. celicara modally have 1
internal scale, the frequency of 0 internasals (=nasals in contact) is much greater
(35%) in S. celicara than in either sample (11% in each) of S. notatus. Throat,
chest, and ventral keeling is summarized in Table 1.

The head pattern of male S. notatus atactus changes with age from a vue
(=female) trilineate pattern to a dotted one and finally to an unpatterned one. Six
males (50%) have head dotting, four (33%) have the head somewhat dotted, and
two (17%) lack head dotting. In Sample 2, 16 males (37%) have the head either
dotted or trilineate, two (5%) have the head somewhat dotted, and 25 (58%) lack
head dotting. Head dotting is present in seven (78%) of nine male S. celicara. In
throat spotting, Sample 1 has six of 12 males (50%) with spotted throats, Sample

VOLUME 95, NUMBER 2 SoM]

2 has nine of 35 males (20%) with the throat spotted, whereas seven of nine (78%)
male S. celicara have the throat spotted.

The female scapular pattern is absent in all 38 females in Sample 1 but occurs
in 18 of 70 females (26%) in Sample 2. In S. celicara, 21 of 22 females (95%) have
scapular patches and ocelli.

Remarks.—We have no doubt that S. celicara is indeed a distinct biological
entity. Whether it would be more appropriate to consider it a subspecies of S.
notatus rather than a separate species is a problem. Sphaerodactylus notatus and
S. celicara are not known to be sympatric, but the ranges of the two approach
fairly closely. The area from which S. celicara is known is remote and difficult
to reach, and the region between the known ranges of S. celicara and S. notatus
remains virtually unknown herpetologically.

Most of the type-series of S. celicara has been taken in or near the Hotel
Asuncion, on the walls and in the lobby. One male was secured in a curujey
(Bromeliaceae) at La Maquina, and another was clinging to arock. Like S. notatus,
S. celicara apparently can adapt easily to edificarian situations, but the species
may occupy a wide variety of niches.

Etymology.—The name celicara is a noun in apposition derived from the Greek
kelis, meaning spot or stain, and kara (an indeclinable neuter noun) meaning
head, in allusion to the spotted heads in both sexes.

Acknowledgments

In addition to specimens in the Instituto de Zoologia (IZ) La Habana, and the
Albert Schwartz Field Series (ASFS), we have examined material in the American
Museum of Natural History (AMNH) and the Museum of Comparative Zoology
(MCZ) through the cooperation of the curators and their assistants: Richard G.
Zweifel, George W. Foley, Ernest E. Williams, and José P. Rosado. The senior
author had assistance in the field from Luis de Armas, Lorenzo Zayas, Jorge de
la Cruz, Luis R. Hernandez, José Espinosa, A. Vega, H. Sarazua, and Miguel
L. Jaume. The senior author also wishes to extend thanks to Fernando Gonzalez,
director of the Instituto de Zoologia, Academia de Ciencias de Cuba, for organ-
izing the trip to Baracoa, and to the authorities of ‘‘Poder Popular’’ for providing
transporation and accommodations in the environs of Baracoa. Jan Krescek sup-
plied us with a transparency of an individual from Monte Iberia. The drawings
of S. celicara are the work of Alvis Gineika, to whom we are also grateful.

(OHG) Instituto de Zoologia, Academia de Ciencias, Capitolio Nacional, La
Habana, Cuba; (AS) Miami-Dade Community College, North Campus, Miami,
Florida 33167.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 398-402

CYRTOCARA LIEMI, A PREVIOUSLY UNDESCRIBED
PAEDOPHAGOUS CICHLID FISH (TELEOSTEI:
CICHLIDAE) FROM LAKE MALAWI, AFRICA

Kenneth R. McKaye and Catherine Mackenzie

Abstract.—A new Lake Malawi paedophagous cichlid, Cyrtocara liemi, is de-
scribed. Cyrtocara liemi is probably most closely related to C. orthognathus.
The ecology and behavior of C. liemi is discussed briefly.

The cichlid species flocks of the Great Lakes of East Africa are the most
spectacular examples of speciation and adaptive radiation in any vertebrate fam-
ily. Each of the lakes, Malawi, Tanganyika and Victoria, contains more species
of fish than any lake outside the Rift Valley region. Lake Malawi has about 300
described species of fish, 90% of which are cichlids, and 100-200 are believed
still to be described (pers. obs.; Marsh et al. 1981). Cichlids are well known for
their great diversity of feeding adaptations (Fryer and Iles 1972) which allows
them to exploit a wide array of foods, including fish eggs, fish embryos and fish
larvae. The existence of paedophagous species, i.e. those which primarily exploit
fish embryos and larvae, was first reported for the Lake Victoria cichlid flock
(Greenwood 1959, 1967, 1974). At least three species of paedophagous cichlids
occur in Lake Malawi, Cyrtocara orthognathus (Trewavas, 1935) and two un-
described cichlids (McKaye and Kocher, in press). This report describes the one of
those paedophagous species of which we have sufficient material. The counts and
measurements employed follow Barel et al. (1977).

Cyrtocara liemi, new species
Fig. 1

Holotype.—National Museum of Natural History (USNM) 227497, adult male,
175 mm standard length (SL) from W. Thumbi Island, Lake Malawi, Malawi
(34°40’E, 14°01’S), collected by K. McKaye, T. Kocher and M. Oliver, Field No.
MKO 80-49, July 1980.

Paratypes.—USNM 227498 (4 specimens, 136.2 mm, 154.1 mm, 166.5 mm,
175.2 mm SL, north side of W. Thumbi Island); 227499 (2 specimens 191.3 mm,
174.2 mm SL, south side of W. Thumbi Island); 227500 (1 specimen, 178.3 mm
SL, Chikale Beach, Nkhata Bay, 34°17’E 11°35’S); 227501 (1 specimen, 124.3 mm
SL, swamp east of Chembe village, Nankumba Peninsula, 34°51’E, 14°01’S);
227502 (1 specimen, 131.8 mm SL); 227503 (1 specimen, 130.4 mm SL, Otter
Point 34°48'E, 14°02’S); 227504 (2 specimens, 60.7 mm, 62.4 mm SL, south east
corner of W. Thumbi Island).

Diagnosis.—Cyrtocara liemi is similar to and probably most closely related to
another paedophagous cichlid C. orthognathus (Fig. 2). The two species can be
easily distinguished by the profile of the head, particularly gape inclination, which
in C. liemi is 40-60° (x = 43.4) but in C. orthognathus is 60-80° (x = 66.6), and

VOLUME 95, NUMBER 2 399

Fig. 1. Holotype of Cyrtocara liemi; USNM 227497.

the premaxillary pedicel inclination, which ranges from 20-30° (x = 23.7) in C.
liemi and 0-10° (x = 1.1) in C. orthognathus. Both species belong to a distinct
group with a single oblique stripe within the genus Cyrtocara and are restricted
to Lake Malawi.

Description.—This description is based on the holotype (Fig. 1) and twelve
- paratypes, ten adults 124.3-191.3 mm SL and two juveniles 60.7 mm and 62.4
mm SL. The principal morphometric ratios are given in Table 1.

The portion of the head anterior to the interorbital region is concave in profile
in larger specimens and nearly straight in smaller specimens. Premaxillary pedicel
inclination 20-—30°. Inclination of the dorsal head profile 28—40°. Snout 1.2-1.6
times longer than broad. Cephalic laterosensory pores and canals not hypertro-
phied.

Posterior margin of maxilla does not extend to the vertical through the anterior
margin of orbit. Jaws narrowly rounded when viewed from above. Lips not no-
ticeably thickened. Lower jaw projects slightly. Gape inclination 40—60°. Lower
jaw 1.15-1.98 times longer than broad.

Fig. 2. Syntype of Cyrtocara orthognathus from British Museum; BM(NH) 1936.6.14.

400 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Principal morphometric ratios of Cyrtocara liemi. % = percent of head length, * =
percent of standard length, M, = mean of adult specimens, M; = mean of juvenile specimens.
R, = range of adult specimens, R, = range of juvenile specimens.

Character M, R, M, R;
Standard length mm 157.9 124.3-191.3 61.6 60.7—62.4
Body depth * 38.5 36.1-42.1 Sil 34.6-39.5
Head length mm 5123 39.5-61.0 | 21.3—22.9
Head length * 3.5) 31.4-33.8 34.1 32.4-35.9
Head breadth % S729 36.4-40.3 40.1 39.3-40.8
Interorbital width % 21.0 1957-22-11 IP).8 17.9-20.7
Snout length % 37.1 34.7-39.1 Sl 28.8-33.3
Orbit length % 24.6 22.5—26.5 29.0 28.2—-29.7
Cheek depth % 19.8 17.0-22.0 1252 12.2—12.2
Post-orbital head length % 40.5 38.2-42.9 36.4 36. 1-36.7
Upper jaw length % 34.4 32.6-36.2 33.5 31.5-35.4
Premaxillary pedicels % 27.0 25.4-30.4 31.5 29332
Lower jaw length % 44.2 42.7-45.9 44.8 43.2-46.3 |
Predorsal length * 33.8 33.1-34.6 36.4 32.3-40.5
Dorsal fin base * 57.6 53.9-59.1 SU sd) 55.5—59.5
Caudal peduncle length * 15.3 NEO = Nes 16.7 16.5-16.8

Caudal peduncle 13.9-16.8% SL. 1.15—1.5 times longer than deep.

Dorsal fin with 27 (1), 28 (10) or 29 (2) elements, comprising 16 (1), 17 (10), 18
(2), spines and 10 (2), 11 (9) or 12 (2) segmented rays; last spine 11.9-15.3% SL.
Anal fin with 10 (1), 12 (3) or 13 (9) elements, comprising 3 spines and 7 (1), 9 (3)
or 10 (9) segmented rays. Pectorals 27.8-32.5% SL comprising 11-14 segmented
rays. Pelvic fins 23.9-31.1% SL, comprising 5 segmented rays. Caudal fin emar-
ginate, lobes pointed, subequal, scaled over entire surface.

Scales ctenoid. Lateral line with 31-34 pored lateral line scales (mode 31),
upper part 23-26 (mode 25), lower part 6—9 (mode 8) Cheek with 3-4 rows (mode
3); uppermost scale often significantly larger. 5 to 7 scales between dorsal-fin
origin and lateral line (mode 5). 7 to 9 scales (mode 8) between pectoral and
pelvic-fin bases, scales gradually decreasing in size from pectoral fin to pelvic fin.
Sixteen scales around caudal peduncle.

Four gill rakers on epibranchial + 1 (in angle) + 10 (1), 11 (11), 12 (1) on cer-
atobranchial. Anterior 4 on ceratobranchial often short, peg-like, remainder lon-
ger, some club-like and bent laterally.

Outer tooth row of upper jaw with a total of 42-60 teeth in adults; two juvenile
specimens with 38 and 39. Outer rows of teeth of both upper and lower jaws
composed of somewhat recurved, slightly movable teeth, embedded in fleshy
gums. Teeth broad, long, and unequally bicuspid anteriorly, becoming shorter,
more slender, and unicuspid posteriorly. Anterior teeth pale, distal portion rusty
brown, color decreasing in extent and intensity in more posterior teeth; posterior-
most teeth uniformly pale. In bicuspid teeth, lateral margin of major cusp convex,
medial margin concave; inner margin of minor cusp more oblique than outer.
Major cusp usually, but not always, anterior to minor cusp. Teeth of larger spec-
imens uniformly unicuspid. Inner tooth rows irregular, 2-3 rows on upper and
lower jaws, teeth unicuspid, occasionally weakly tricuspid with middle cusp ex-
tended.

VOLUME 95, NUMBER 2 401

Fig. 3. Lower pharyngeal bone of Cyrtocara liemi in occlusal view.

Lower pharyngeal bone (n = 7) (Fig. 3) triangular in outline, 1.13—1.5 times
broader than long, its length 10.9-28.4% of head length, breadth 57.1-72.7% of
head breadth. Suture between two halves straight. Dentigerous area 1.2—1.6 times
broader than long. Teeth variable. Postero-median teeth large but decreasing in
size toward end of row, weakly bicuspid, with posterior cusp longest and colored
rusty brown at tip and curved forward slightly. Lateral teeth compressed, bicus-
pid or unicuspid, paler in color. Teeth closely packed posterolaterally, more
openly packed medially, 29-35 teeth in posterior row, 5—11 in median row, 4-8
in oblique rows, and 19-26 in lateral rows.

Coloration in preservation.—Both sexes with head and dorsal surface dark
grey-brown, paling posteriorly and ventrally. Sides with single oblique stripe,
divided into 3 overlapping sections. Anterior section extending posteriorly to
scale 13-18 of lateral line, beginning above the upper part of the lateral line and
crossing over it at scale 7-8. Middle section extending from scale 7-9 to scale
19-28, between upper and lower lateral line sections. Posterior section beginning
at scale 16-19 and extending along upper side of lower lateral line to base of
caudal fin where it dips slightly below the lower lateral line. Fins translucent
brown-grey, with pale maculae. Juveniles pale yellow-brown. Oblique stripe com-
posed of discontinuous segments. 10 vertical bars distinguishable. Fins colorless
to grey, translucent.

Distribution.—This species is not common, but it does occur in the Cape Ma-
clear (Nankumba Peninsula) region of Lake Malawi as well as in Nkhata Bay. It

402 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

was caught and observed by SCUBA divers (McKaye and Kocher, in press) only
in areas where brooding cichlid females congregated.

Relationships.—An analysis of the phylogenetic relationship of C. liemi must
await revision of the genus Cyrtocara, which includes the group of obliquely-
striped species of which the paedophages C. liemi and C. orthognathus are mem-
bers. These fishes with a single oblique stripe have been grouped by Trewavas
(1935) in her key and may represent a monophyletic group. Cyrtocara liemi and
C. orthognathus occur sympatrically at Cape Maclear and at Nkhata Bay. Not
only do they differ significantly in morphology, they also differ in behavior.

Etymology.—Named after Karel Liem in recognition of his pioneering studies
and his insight into the feeding mechanism of cichlid fishes.

Ecology.—Cyrtocara liemi is a paedophagous cichlid which acquires eggs, em-
bryos, and fry from the mouths of mouthbrooding cichlids. Cyrtocara liemi at-
tacks females from 0.5 to 2 m below and hits them in the hyoid region of the
head. The only items found in their stomachs were eggs, embryos and larval
cichlids. A more detailed description of their behavior and ecology along with
two other paedophagous cichlids is discussed elsewhere (McKaye and Kocher,
in press).

Acknowledgments

We thank L. Knapp and R. Vari of the Smithsonian Institution for their en-
couragement and guidance, and T. Kocher, D. Lewis and M. Oliver for discussion
of the material. The junior author wishes to thank L. Buss for his generosity and
support throughout this project. We especially thank the government of Malawi
for providing the facilities to make this research possible, and the Cape Maclear
fisheries technicians, R. D. Makwinje, W. M. Menyani, and O. K. Mhone for
help in making our collections. Financial support was from the National Science
Foundation DEB 79-12338, Smithsonian Sorting Center, World Wildlife Fund
U.S.A., and Peabody Museum, Yale University.

Literature Cited

Barel, C. D. N., M. J. P. van Oijen, F. Witte, and E. L. M. Witte-Maas. 1977. An introduction to
the taxonomy and morphology of the haplochromine Cichlidae from Lake Victoria.—Nether-
land Journal of Zoology 27(4):339-389.

Fryer, G., and T. D. Iles. 1972. The cichlid fishes of the Great Lakes of Africa.—Oliver & Boyd,
Edinburg, 641 pp.

Greenwood, P. H. 1959. A revision of the Lake Victoria Haplochromis species (Pisces, Cichlidae),

Part 3.—Bulletin British Museum of Natural History (Zoology) 5:179-218.

. 1967. A revision of the Lake Victoria Haplochromis species (Pisces, Cichlidae), Part 6.—

Bulletin British Museum of Natural History (Zoology) 25:139-242.

. 1974. The cichlid fishes of Lake Victoria, East Africa: The biology and evolution of a species

flock.—Bulletin British Museum of Natural History (Zoology) Supplement 6: 1-134.

Marsh, A. C., A. J. Ribbink, and B. A. Marsh. 1981. Sibling species complexes in sympatric pop-
ulations of Petrotilapia Trewavas (Cichlidae, Lake Malawi).—Journal Linnean Society (Zo-
ology) 71:253-—264.

McKaye, K. R., and T. Kocher. Head ramming behavior in three paedophagous cichlids in Lake

Malawi, Africa.—Animal Behavior (in press).
Trewavas, E. 1935. A synopsis of the cichlid fishes of Lake Nyasa.—Annals and Magazine of Natural

History 16:65-118.

(KRM) Duke University Marine Laboratory, Beaufort, North Carolina 28516,
and (CM) Ecologia, I.N.P.A. EP 478, Manaus 69000, Brazil.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 403-407

HYPSELOGNATHUS HORRIDUS, A NEW SPECIES OF
PIPEFISH (SYNGNATHIDAE) FROM SOUTH AUSTRALIA

C. E. Dawson and C. J. M. Glover

Abstract.—An unusual pipefish, characterized by spinigerous head and body
surfaces, absence of scutella, and presence of caudal fin and caudal brood pouch,
is described from depths of 42-55 m off Anxious Bay, South Australia. Provi-
sionally referred to the Australian endemic genus Hypselognathus, this species
differs from its only congener (H. rostratus) in having spiny rather than essentially
smooth head and body surfaces, in lacking scutella, in having more total subdorsal
rings (11-11.5 versus 8—9.5) and a higher HL in SL ratio (8.0—9.0 versus 5.9-6.6),
as well as in other morphological, proportional and meristic features.

Current studies on Australian pipefishes indicate the presence of about 90 valid
species and several yet undescribed forms. Most of the latter have similar or
closely related congeners and descriptions are delayed pending collection of more
_ comparative material. Specimens reported here differ so strikingly from all known
pipefishes that we see no need for postponing their description. The occurrence
of this species and the recently described Kimblaeus bassensis Dawson (1980) in
moderate depths (40-70 m) of southern Australian coasts suggests that continued
sampling in this region may educe further additions to the pipefish fauna of Aus-
tralia.

Measurements are, in part, referred to standard length (SL) and head length
(HL); counts of the holotype are marked *; as employed here, the term ‘‘venter”’
refers to the ventral surface of head or body; other methods are those of Dawson
(1977). Specimens are deposited in the South Australian Museum (SAM) and Gulf
Coast Research Laboratory Museum (GCRL). Drawings are by Mrs. Yasue Mat-
thews (GCRL).

Hypselognathus horridus, new species
Figs. 1 and 2

Holotype.—SAM F.4676 (154.0 mm SL, adult male), South Australia, Great
Australian Bight, off Anxious Bay, 54.9 m (30 fm.), trawl, 25 Feb. 1981, P. C.
Halsey.

Paratypes.—GCRL 18057 (226.5 mm SL, adult female), South Australia, Great
Australian Bight, ca. 32°24’S, 133°30’E, 42 m, trawl, 5 May 1973, SA Fish. Dept.
SAM F.4681 (228.0 mm SL, adult female), South Australia, Great Australian
Bight, ca. 33°24’S, 134°37’E, 14.8 km S of Point Weyland, 54.9 m, trawl, 18 Sept.
1981, Karl Olsen.

Diagnosis.—Head and body surfaces (except membranes) spinigerous, scutella
absent, HL 8.0-9.0 in SL, snout length 1.6-1.8 in HL, total rings 70-72.

Description.—Rings 27* + 43-45*, subdorsal rings 6.75-6.0* + 4.25-5.5* =
11.0-11.5*, dorsal-fin rays 31*, pectoral-fin rays 10* (2), 11 (2), 12 (2), anal-fin
rays 3*, caudal-fin rays 10*. Measurements of holotype are followed (in paren-

404 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Sate Pa: ~
6 Ooh RE aaa eas
OR a tea oC TNS Se =
Sie Fame Sati 2 eS ee

Fz cuumeg SHY SES TG
< [OE EES. LP pe roeay gs Poe Ga oe

We fe, yg
Lh pee
VV 71.
LTT fi?
Wo
, eek Mf

Fig. 1. Hypselognathus horridus. Upper pair: Lateral and dorsal aspects of head and anterior
trunk rings. Lower pair, Top: Posterior trunk and anterior tail rings, together with dorsal and anal
fins and anterior portion of brood pouch. Bottom: Posterior tail rings and caudal fin. From holotype
(SAM F.4676).

theses) by those of shorter and longer paratypes, respectively: SL 154.0 (226.5,
228), HL 17.1 (28.5, 28.4), snout length 9.3 (17.3, 17.3), least snout depth 2.1
(3.4, 3.2), length of dorsal-fin base 16.4 (24.3, 24.9), anal ring depth 2.6 (4.5, 4.2),
trunk depth 3.2 (6.8, —), pectoral-fin length 3.2 (4.2, —), length of pectoral-fin
base 1.6 (2.9, 2.8), caudal-fin length 7.3 (—, 11.1).

Median dorsal snout ridge (Fig. 1) low, irregularly denticulate to spinulose,
ending just before vertical through anterior margin of orbit, not confluent with
anterior continuations of supraorbital ridges; snout compressed laterally, its least
depth 4.4—5.4 in snout length; nares 2-pored bilaterally, the anterior pore on a

VOLUME 95, NUMBER 2 405

Fig. 2. Hypselognathus horridus. Top and middle: SAM F.4676 (154 mm SL, male, holotype).
Bottom: GCRL 18057 (226.5 mm SL, female, paratype).

short tubule; preorbital bone rather narrow, its width less than diameter of pupil.
Opercle with radiating denticulate striae, none clearly enlarged or ridge-like; dor-
sal margins of orbits elevated slightly; interorbital essentially flat, a little de-
pressed between dorsal margins of orbits; dorsum of head more or less rounded
behind orbits; nuchal ridge obsolete; prenuchal and frontal ridges vestigial; pec-
toral-fin base protruding a little laterad, without obvious ridges.

Superior trunk and tail ridges discontinuous near rear of dorsal-fin base; lateral
trunk and tail ridges discontinuous, terminating on last trunk ring; inferior trunk
and tail ridges continuous (Fig. 1). Principal body ridges distinct, somewhat el-
evated and angled a little laterad on posterior 16-20 tail rings; ridge margins
irregularly denticulate, usually with a slightly enlarged hook-like spine on pos-
terior third of each ring; anterior and posterior margins of rings straight to irreg-
ularly emarginate; scutella absent. Most surfaces (except eye, gill membranes,
fins and pouch folds) more or less covered with minute spines, their distribution
irregular but numbers somewhat reduced on posterior tail rings. Dorsum a little
convex on trunk and anterior half of tail, gradually becoming flat and depressed
between somewhat elevated superior ridges on posterior tail rings; venter of trunk
V-shaped, the median ridge not clearly enlarged; 12th—19th trunk rings of holo-
type swollen or enlarged in dorsal and lateral aspects (Fig. 2); 11th—27th trunk
rings of adult females deeper (ca. 22%) than preceding rings; sides and venter of
posterior tail rings more or less flat, depressed distally between elevated principal
ridges.

406 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Dorsal-fin base not elevated; superior trunk ridge not arched strongly dorsad
below dorsal-fin base; pectoral fin more or less rounded; caudal-fin membranes
broad above and below, narrower between the long median fin-rays; tips of cau-
dal-fin rays somewhat flattened and spatulate.

Brood pouch developed below the anterior 18 tail rings of the holotype; pouch
plates moderately enlarged, angled somewhat laterad; membranous pouch folds
present. Pouch closure, uncertain due to absence of eggs or larvae, probably the
everted type of Herald (1959).

Color of holotype in alcohol mainly grayish, shading to brown on distal half of
tail. Sides and dorsum of head and body, and venter of tail behind brood pouch,
peppered with brown to black microchromatophores, somewhat larger on sides
and most abundant on posterior half of tail; anterior and posterior margins of
rings often with fine dark edging. Lower portions of gill membranes, gular mem-
brane, and venter of trunk mainly pale. Brood-pouch folds pale, with moderately
broad, irregular, brownish shading along mesial margins. Caudal-fin rays largely
brownish, the membranes shaded diffusely with brown; other fins hyaline. Col-
oration similar in a recently preserved female paratype (SAM F.4681) but the
venter of the deepened posterior portion of the trunk is shaded with pink.

Etymology.—Named horridus, from the Latin, meaning bristly or rough.

Relationships.—The combination of spinigerous surfaces, absence of scutella,
presence of caudal fin and a brood pouch, with plates and folds, located under
the tail distinguishes the species described here from all known pipefishes. Its
relationships are unclear and it may eventually warrant separate generic treat-
ment. In general morphology, this fish appears most closely related to the poorly
defined Hypselognathus Whitley (1948) and it is provisionally referred to this
otherwise monotypic, Australian endemic, genus (type species Histiogamphelus
rostratus Waite and Hale, 1921) until additional material is available for study.

Comparisons.—Hypselognathus rostratus and H. horridus agree in the config-
uration of principal body ridges (Fig. 1), in the presence of dorsal, pectoral, anal
and 10-rayed caudal fins, and both share an elongate, laterally compressed, snout.
In general agreement with H. horridus, specimens of H. rostratus may have a
small spine or vestige thereof on margins of principal ridges of trunk rings, prin-
cipal ridges of the posterior tail rings are somewhat elevated and angled laterad,
and some swollen trunk rings are typically present. However, H. horridus lacks
the small round scutella present in H. rostratus, fails to have the supraorbital
ridges confluent with the median dorsal snout ridge (confluent in rostratus), and
has spinigerous head and inter-ridge body surfaces (smooth in rostratus). Com-
pared to 8 specimens of H. rostratus (152-305 mm SL), H. horridus has more
trunk rings and total rings (respectively, 27 and 70—72 versus 24—25 and 65-69),
fewer pectoral-fin rays (10-12 versus 12-14), more subdorsal trunk rings and total
subdorsal rings (respectively, 6.75—6 and 11—11.5 versus 2.5—0.75 and 8-9.5), a
higher HL in SL ratio (8.0-9.0 versus 5.9-6.6), and a lower snout depth in snout
length ratio (4.4—5.4 versus 6.2—13.0).

Among other pipefishes, spinigerous head and body surfaces, somewhat similar
to those of H. horridus, occur only in some species of Solegnathus Swainson
(e.g., S. spinosissimus Gtinther). However, species of Solegnathus differ from
Aypselognathus horridus in lacking the caudal fin and brood-pouch plates and
folds and in having well-developed scutella.

VOLUME 95, NUMBER 2 407

Remarks.—Each of the three known specimens of Hypselognathus horridus
was taken with one or more specimens of Solegnathus robustus McCulloch.

Literature Cited

Dawson, C. E. 1977. Review of the pipefish genus Corythoichthys with description of three new

species.—Copeia 1977:295-338.

. 1980. Kimblaeus, a new pipefish genus (Syngnathiformes: Syngnathidae) from Australia, with

a key to genera of pipefishes with continuous superior ridges.—Australian Journal of Marine

and Freshwater Research 31:517—523.

Herald, E. S. 1959. From pipefish to seahorse—a study of phylogenetic relationships.—Proceedings
of the California Academy of Sciences 42:181—227.

Waite, E. R., and H. M. Hale. 1921. Review of the lophobranchiate fishes (Pipe-fishes and Sea-
horses) of South Australia.—Records of the South Australian Museum 1:293-324.

Whitley, G. P. 1948. Studies in ichthyology. No. 13.—Records of the Australian Museum 22:70-94.

(CED) Gulf Coast Research Laboratory Museum, Ocean Springs, Mississippi
39564; (CJMG) South Australian Museum, Adelaide, S.A. 5000, Australia.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 408-411

LATERAL LINE OF DIPLOSPINUS MULTISTRIATUS
(TELEOSTEI: GEMPYLIDAE)

Izumi Nakamura

Abstract.—The existence of the lateral line in Diplospinus multistriatus has
been questioned. The lateral line of D. multistriatus is described here and com-
pared with that of closely related Paradiplospinus gracilis. Most of the (upper)
lateral line of both species runs nearer the ventral contour of the body than the
dorsal contour. This type of lateral-line conformation is found in some trichiurid
genera, but not in other gempylid genera. Lateral-line scales of both species are
tube-shaped with an irregular longitudinal slit. This type of lateral-line scale is
not found in any other genera of Gempylidae or Trichiuridae.

Since Diplospinus multistriatus was described by Maul (1948) from Madeira,
considerable numbers of specimens have been collected widely in tropical and
temperate waters of the Atlantic, Pacific and Indian oceans (Parin and Bekker
1972:161, Fig. 17). Maul (1948) studied the holotype with six paratypes and wrote,
‘‘the skin is smooth and there is no trace of a lateral line.’’ Later, Tucker (1956)
examined one of the paratypes and reported, ‘“‘there are traces of an apparent
and highly probable lateral line,’’ but he did not mention this further. No other
authors have paid special attention to the lateral line of D. multistriatus.

While examining specimens of D. multistriatus at the National Museum of
Natural History (USNM), Smithsonian Institution, traces of a lateral line were
recognized. As there are no undamaged specimens of either D. multistriatus or
Paradiplospinus gracilis, these observations are based on many specimens com-
pounded to reconstruct the lateral line.

Material examined.—Diplospinus multistriatus Maul: USNM 215428, 2 speci-
mens (16.5, 55.2 mm SL), 21°30’N, 158°30’W, 19 Jan. 1970; USNM 215449, 6
(42.1-188.4), 21°30’N, 158°30’W, 23 Sep. 1970; USNM 100492, 1 (48.0), 29°00’N,
76°23'W, 28 Feb. 1914; USNM 226993, 3 (98.2-162.6), 27°45’N, 91°18.5'W, 23
Feb. 1964; USNM 215397-8, 3 (134.7-203.0), 21°30'N, 158°30'W, 28 Feb. 1971;
USNM 215395, 1 (141.0), 21°30’N, 158°30’W, 17 Sep. 1970; USNM 215433, 2
(142.5, 152.4), 21°30’N, 158°30’'W, 16 Sep. 1970; USNM 194458, 1 (175.1),
29°40’N, 69°05’'W, 29 Mar. 1957; USNM 219967, 2 (175.6, 213.3), 33°04'N,
39°29'W, 27 Apr. 1979; USNM 215391, 1 (188.4), 21°30’N, 158°30’W, 15 Sep.
1970; USNM 215394, 1 (203.8), 21°30’N, 158°30’W, 14 Dec. 1970; USNM 215454,
1 (232.8), 21°30’N, 158°30’W, 18 Sep. 1971.

Paradiplospinus gracilis (Brauer): USNM 208104, 1 specimen (47.7 mm SL),
49°06'S, 120°15’W, 19 Dec. 1965; USNM 208446, 9 (141.0-335.0), 40°18’S,
39°04’W, 8 Mar. 1971; USNM 208448, 6 (161.7—320.0), 39°47’S, 43°38’W, 7 Mar.
1971; USNM 208449, 9 (182.0-349.0), 38°20’S, 54°33’W, 5 Mar. 1971; USNM
226992, 1 (355.0), 40°08'S, 82°47'W, 2 Oct. 1966.

Identification

Diplospinus multistriatus and Paradiplospinus gracilis are similar in external
appearance. Specimens obtained by usual collecting methods are almost always

VOLUME 95, NUMBER 2 409

Table 1.—Some distinguishing characters (counts compared with those of some authors) of Diplo-
spinus multistriatus and Paradiplospinus gracilis.

Characters Species Diplospinus multistriatus Paradiplospinus gracilis

Position of anus midway between tip of snout and tip __ nearer tip of caudal fin than to tip of

of caudal fin; in front of first anal snout; in front of first anal spine by
spine by distance equal to head distance equal to snout length (Fig.
length (Fig. 1A) 1B)
Anal fin anterior part very low, with almost no anterior part fairly high, with fin
fin membrane (Fig. 1A) membrane (Fig. 1B)
Lateral line easily removed; double ? (Fig. 1A) usually intact; single (Fig. 1B)
Dorsal-fin rays XXXI-XXXIV, 35—40 (this study) XXX VI-XXXIX, 28-33 (this study)
XXX-XXXIV, 36-42 (Parin and XXX VI-XXXIX, 28-33 (Parin and
Bekker 1972) Bekker 1972)

XXXI-XXXIII, 37-41 (Karrer 1975) XXXVI-XXXVII, 28-30 (Karrer 1975)
XXXU-XXXVI, 37-41 (Parin et al. XXX VI-XXXVII, 28-32 (Bussing

1978) 1965)
XXXII-XXXIII, 40 (Tucker 1956) XXXVI-XXXVII, 28-32 (DeWitt and
Hureau 1979)
Anal-fin rays II, 28-33 (this study) II, 26-31 (this study)
II, 29-32 (Parin and Bekker 1972) II, 25-30 (Parin and Bekker 1972)
II, 28-34 (Karrer 1975) II, 26-28 (Karrer 1975)
II, 28-33 (Parin et al. 1978) II, 25-28 (Bussing 1965)

II, 25-28 (DeWitt and Hureau 1979)

Vertebrae number 22-24 + 34-37 = 57-61 (this study) 32-34 + 32-34 = 65-66 (this study)
59-61 (Karrer 1975) 61 (Karrer 1975)
34 + 24 = 58 (Tucker 1956) 63-67 (DeWitt and Hureau 1979)
63-66 (Bussing 1965)
38-40 + 26-27 = 65-66 (Andriashev
1960)

more or less damaged and the vertical-fin rays are folded with damaged fin mem-
branes. Some distinguishing characters of the species are summarized in Table
1. Useful characters for distinguishing the species are the number of dorsal spines
and the number of dorsal soft rays (the total number of both is not useful) and
the position of the anus. The number of anal-fin rays overlaps, so it is not a very
useful character. The shapes of the anal fin and the lateral line are often hard to
see. The number of vertebrae (total) clearly differentiates the species.

Lateral Line

The lateral line of D. multistriatus is easily removed, but careful examination
with a microscope can find remains somewhere on the bodies of even fairly
damaged specimens. The remains are found most often on the shoulder region
and next most often on the mid-portion of the body. The lateral-line system based
on several specimens compounded is shown in Fig. 1A. The upper lateral line is
recognized certainly. The lower lateral line was discerned only partly in three out
of 24 specimens examined, and it is not certain whether the lower lateral line
connects with the upper lateral line or where its anterior and posterior termina-
tions may be. From the upper margin of the opercle the upper lateral line of D.
multistriatus descends gently to below the middle of the body at the anus, there-
after running near the ventral contour of the body to the caudal region. The shape

410 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. A, A reconstruction of the lateral-line system of Diplospinus multistriatus based on several
specimens; B, A reconstruction of the lateral-line system of Paradiplospinus gracilis based on several
specimens. Vertical broken lines show the position of the anus. a-g shows the details of lateral-line
scales, a—d from the positions shown in D. multistriatus (A) and e-g from P. gracilis (B). a-c: USNM
215454, 232.8 mm SL; d: USNM 215428, 55.2 mm SL; e: USNM 208446, 255.5 mm SL; f: USNM
226992, 355.0 mm SL; g: USNM 208449, 190.5 mm SL. Scales indicate 1 mm.

of the lateral-line scales is slightly different in various parts of the body as shown
in Fig. la—d. Each scale is basically tube-shaped with an irregular longitudinal
external slit.

The lateral line of P. gracilis is conspicuous, tough, and not usually lost. Most
of the lateral line remains intact in even fairly damaged specimens. The lateral-
line system, based on several specimens, is shown in Fig. 1B. The lateral line of
P. gracilis is single, descending gently from the upper margin of the opercle to
slightly below the middle of the body at the anus, thereafter running slightly
nearer the ventral contour of the body to the caudal region (Fig. 1B). The shape
of the lateral-line scales is somewhat different in various parts of the body, as
shown in Fig. le-g. Each scale is basically a short tube with an irregular external
slit. Scales in the middle of the lateral line (Fig. le) are much more elongate than
those in the anterior and posterior parts (Fig. 1f, g). Lateral-line scales are gen-
erally stouter in P. gracilis than in D. multistriatus. |

The course of the (upper) lateral line is similar in both species. Careful obser-
vation, however, reveals that most of the (upper) lateral line is situated only
slightly below mid-body in P. gracilis and far below mid-body in D. multistriatus.
Excluding the lower lateral line of D. multistriatus from consideration, similar
lateral lines running below mid-body are found in some trichiurid genera, such
as Trichiurus, Lepturacanthus, Eupleurogrammus and Tentoriceps, but not in
any other gempylid genera, which have the single lateral line or the upper part
of a double lateral line running nearer to the dorsal countour than the ventral
contour. This may suggest a close relationship of gempylids and trichiurids.

VOLUME 95, NUMBER 2 411

The basic structure of the lateral-line scales of D. multistriatus and P. gracilis
are similar. This type of lateral-line scale (tube-shaped scale with an irregular
longitudinal external slit which has relatively small openings at both ends of each
scale) is not found in any other genera of Gempylidae and Trichiuridae.

The lateral line seems to become fully formed at about 50 mm SL in both D.
multistriatus and P. gracilis. The size at which the lateral-line development starts
could not be ascertained, though some larval materials of both species were
examined in this study. Specimens of D. multistriatus about 20 mm SL do not
show any traces of the lateral line (Strasburg 1964; Yevseyenko and Serebryakov
1973) and a specimen of P. gracilis 32.2 mm SL does not show any traces of it
(Bussing 1965).

Acknowledgments

I am grateful to the Smithsonian Institution for providing me a postdoctoral
fellowship in 1981, which made it possible for me to do this study. Thanks are
also due to my supervisor, Robert H. Gibbs, Jr. and Joseph L. Russo of NMFS
Systematics Laboratory for their critical review. Technical assistance rendered
by Reiko Nakamura is acknowledged.

Literature Cited

Andriashev, A. P. 1960. Families of fishes new to the Antarctic. I. Paradiplospinus antarcticus gen.
et sp. (Pisces, Trichiuridae).—Zoologicheskii Zhurnal 39(2):244—249 (in Russian).

Bussing, W. A. 1965. Studids of the midwater fishes of the Peru-Chile Trench.—American Geo-
physical Union, Antarctic Research Series 5:185—227.

DeWitt, H. H., and J.-C. Hureau. 1979. Fishes collected during *‘Hero’’ Cruise 72-2 in the Palmer
Archipelago, Antarctica, with the description of two new genera and three new species.—
Bulletin du Muséum National d’ Histoire Naturelle, Paris 4° série 1:775-820.

Karrer, C. 1975. Uber Fische aus dem Siidostatlantik (Teil 2).—Mitteilungen aus dem Zoologischen
Museum in Berlin 51(1):63-82.

Maul, G. E. 1948. Quatro peixes novos dos mares da Madeira.—Boletim do Museum Municipal do
Funchal (3) Art. 6:41-55 (in Portuguese and English).

Parin, N. V., and V. E. Bekker. 1972. Materials on taxonomy and distribution of some trichiuroid

fishes (Pisces, Trichiuroidae: Scombrolabracidae, Gempylidae, Trichiuridae).—Transactions

of the P. P. Shirshov Institute of Oceanography 93:110-—204 (in Russian).

, Y. I. Sazonov, and S. V. Mikhailin. 1978. Glubokovodnye Pelagicheskie Ryby V Clorkh Ips

‘‘Fiolent’? V Gvineiskom Zalive i Prilegayoshchikh Raionakh.—Trudy Instituta Okeanologii

im P. P. Shirshova 3:169—183 (Translated from Russian for the Smithsonian Institution and the

National Science Foundation, Washington, D. C. by Esduck Cairo: Deep-water pelagic fishes

in the collections of R/V ‘‘Fiolent’’ in the Gulf of Guinea and adjacent regions. 1981).

Strasburg, D. W. 1964. Postlarval scombroid fishes of the genera Acanthocybium, Nealotus, and
Diplospinus from the Central Pacific Ocean.—Pacific Science 18(2):174—-185.

Tucker, D. W. 1956. Studies on the trichiuroid fishes—3. A preliminary revision of the family Trichi-
uridae.—Bulletin of the British Museum (Natural History) Zoology 4(3):73-130.

Yevseyenko, S. A., and V. P. Serebryakov. 1973. Larvae of Diplospinus multistriatus (Pisces, Gem-
pylidae) from the northwestern Atlantic.—Journal of Ichthyology 14(1):92-98.

Fisheries Research Station, Kyoto University, Maizuru, Kyoto 625, Japan, and
Division of Fishes, Department of Vertebrate Zoology, National Museum of Nat-
ural History, Smithsonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
95(2), 1982, pp. 412-420

VARIATION OF A MUSCLE IN HUMMINGBIRDS AND
SWIFTS AND ITS SYSTEMATIC IMPLICATIONS

Richard L. Zusi and Gregory Dean Bentz

Abstract.—Historically, features of the muscle tensor propatagialis pars brevis
have been used to argue that swifts and hummingbirds comprise a single mono-
phyletic order, the Apodiformes, and that this order is most closely related to the
‘‘pico-passeriforms.’’ We describe variations of this muscle in swifts and hum-
mingbirds as well as in other orders and conclude that the new evidence does not
support these claims. The variations observed, however, do show morphological
trends that help to clarify relationships within both swifts and hummingbirds.

The phylogenetic relationships of hummingbirds and swifts to each other and
to other avian orders are among the major unsolved ornithological problems in
systematics. Subfamilial and generic relationships within swifts are fairly well
understood (see Brooke 1970), but internal relationships of hummingbirds are still
poorly known. Although we do not claim to have solved any of these problems,
we have found that variation in a single muscle, M. tensor patagii brevis, bears
on all of them. We report this variation and its systematic implications as a
stimulus for further study of these problems, and as partial evidence toward their
eventual solution.

The tensor patagii brevis muscle (TPB) extends from the shoulder to the fore-
arm in birds and apparently can serve either to flex the forearm or to support the
prepatagial membrane of the extended wing. This muscle exhibits marked vari-
ation among birds, especially in relation to its tendon or tendons of insertion.
Garrod (1876) drew attention to this variation and to its taxonomic implications,
and Fiirbringer (1888) devoted six plates and considerable discussion to this mus-
cle. Since then it has been further described in major systematic and anatomical
works (e.g. Buri 1900; Beddard 1898; George and Berger 1966). In this paper we
describe in some detail the variation of TPB in swifts and hummingbirds, and we
test Lowe’s (1939:329) contentions that the fleshy belly and tendon of insertion
of TPB in hummingbirds is almost identical with that of the swift, that both are
fundamentally passerine in design, and that the arrangement in non-passerines is
quite different.

Hummingbirds

In hummingbirds the origin of TPB is consistently by a tendon from the head
of the coracoid. The nearly parallel fibers of the short, wide belly pass distally
and end on the surface of the extensor metacarpi radialis muscle (EMR) and on
a short internal aponeurosis that fuses with the aponeurosis of origin of EMR. At
this point of fusion the aponeurosis of TPB forms a tendon (the humeral tendon)
that extends across the belly of EMR and inserts on the humerus. In humming-
birds another aponeurosis or tendon (the distal tendon) passes distally from the

VOLUME 95, NUMBER 2 413

same point of fusion along the surface of EMR toward the wrist. Within hum-
mingbirds, four different patterns of insertion exist.

Type 1 (Fig. 1f) is characteristic of the Phaethornithinae or hermits (see Ap-
pendix for species examined). In these forms the humeral tendon of TPB is par-
tially covered by fibers of EMR that arise from the process of origin of that
muscle. The humeral tendon of TPB and the aponeurosis of origin of EMR are
tightly bound together where they cross within the belly of EMR, thus forming
a firm base for the attachment of overlying muscle fibers. The humeral tendon of
TPB emerges from the belly of EMR, passes superficial to the distal-most fibers
of origin of EMR and inserts on a tubercle of the humerus distal to the process
of EMR. (Such terms as ectepicondylar process, lateral epicondyle, and dorsal
supracondylar process are sometimes used for the points of attachment of EMR
and TPB on the humerus. Until a better understanding of the homologies of
projections on the distal end of the humerus is obtained throughout birds we
prefer to avoid these terms and refer simply to the process of EMR and the
tubercle of TPB.)

The distal “‘tendon’’ of TPB in hermits is so transparent that we initially con-
sidered it absent. Closer examination revealed a short tendon from the junction
of the bellies of TPB and EMR that quickly fans out into a broad aponeurotic
sheet and invests most of the distal half of the belly of EMR. Proximally it is free

of a superficial dorsal aponeurosis of EMR, but distally the two aponeuroses fuse.
_ Eutoxeres is the only phaethornithine exception to the Tyoe 1 format. In this
genus the humeral tendon of TPB is visible for its entire length. Although not
covered by fibers of EMR, it serves as a point of origin for some of the superficial
fibers of that muscle. The humeral tendon and the aponeurosis of origin of EMR
are fused where they cross as in other hermits.

Type 2 (Fig. 1g) occurs in some of the Trochilinae: Anthracothorax, Doryfera,
Androdon, Eulampis, Sericotes, Chrysolampis, Polytmus, Topaza, Heliothryx,
Colibri, and Florisuga. In these forms the humeral tendon of TPB is clearly
visible on the surface of the proximal end of EMR. Unlike Type 1 the humeral
tendon is essentially free of the aponeurosis of origin of EMR, attaching only to
its cranial edge. The insertion of the humeral tendon is on a tubercle of the
humerus. As in other trochiline hummingbirds a distinct distal tendon of TPB is
formed. This tendon broadens distally and fuses with the dorsal surface of a
superficial aponeurosis of EMR that contributes to its tendon of insertion near
the wrist. This distal tendon of TPB lies on the dorsal surface of EMR so that the
tendon appears to divide the belly of EMR unequally into cranial (smaller) and
caudal (larger) portions. This is not the case with Type 3 birds.

Type 3 (Fig. 1h) is found in all trochiline genera listed in the Appendix except
those of Types 2 and 4. The distal tendon of TPB is well developed and, at least
proximally, free of the belly of EMR. It passes along the cranial edge of the belly
of EMR rather than along the dorsal surface. Only near the distal end of the
radius does it fuse with the craniodorsal portion of the tendon of insertion of
EMR. The relations of the humeral tendon of TPB are like Type 2. This is the
type illustrated by Garrod (1876).

Type 4 (Fig. 11) is restricted to trochiline hummingbirds of the genera Aces-
trura, Chaetocercus, Calliphlox, and possibly others. Here the relations of the

414 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Diagrams showing M. tensor propatagialis pars brevis and M. extensor metacarpi
radialis (right, dorsal view): a, Piciformes and some Passeriformes; b, Most Passeriformes (dotted
outline represents Hirundinidae and Lonchurae); c, Hemiprocnidae (Type A); d, Cypseloidinae (Type
B); e, Apodinae (Type C); f, Trochilidae (Type 1); g, Trochilidae (Type 2); h, Trochilidae (Type 3); i,
Trochilidae (Type 4). EMR = M. extensor metacarpi radialis; PPB = M. pectoralis pars propatagialis
brevis; TPB = M. tensor propatagialis pars brevis. Long arrow indicates humeral tendon of TPB; short
arrow indicates distal tendon of TPB.

distal tendon of TPB are similar to Type 3, but a difference exists in the humeral
tendon. In these forms, that tendon lies directly superficial to and almost parallel
with the aponeurosis of origin of EMR, to which it is firmly fused. The short
tendon of TPB inserts on the process of origin of EMR rather than on a separate
tubercle.

VOLUME 95, NUMBER 2 415

Swifts

The condition of TPB in swifts is significantly different from that of humming-
birds. According to Cohn (1968) the muscle arises from the head of the coracoid
and from the adjacent dorsal arm of the furcula in larger swifts, and only from
the coracoid in some smaller swifts. Furthermore, M. pectoralis pars propatagialis
longus and M. pectoralis pars propatagialis brevis are present in swifts but neither
is present in hummingbirds. In swifts the latter muscle is represented by a short
tendon that extends from M. pectoralis near the deltoid crest to the tendon of
insertion of TPB. In those instances where the tendon of insertion of TPB is
absent, the propatagialis brevis joins TPB at its juncture with EMR. In all of the
swifts we examined there was no trace of a distal tendon of TPB. Within the
swifts the TPB differs in other ways as well, and we recognize three basic types.

Type A (Fig. 1c) is found in the crested swifts (Hemiprocnidae). In these forms
the belly of TPB ends on a short tendon that receives the pectoralis pars pro-
patagialis brevis tendon and continues to the cranial surface of EMR. The humeral
tendon then passes across the surface of EMR to insert on a tubercle of the
humerus.

Type B (Fig. 1d) is found in the Cypseloidinae. It is similar to Type A except
that the belly of TPB reaches the surface of EMR without first forming a tendon
of insertion. Instead, the belly tapers to a narrow, semitendinous insertion on
EMR. At this juncture pars propatagialis brevis attaches and a well-defined hu-
meral tendon of TPB arises and crosses EMR to insert on a tubercle of the
humerus.

Type C (Fig. le) is seen in the Apodinae (sensu Morony et al. 1975; includes
Collocaliini, Chaeturini, and Apodini). Type C resembles Type B except that the
belly of TPB makes broad contact with EMR rather than tapering to its insertion.
In addition, the humeral tendon of TPB almost parallels that of EMR and inserts
directly on the process of origin of EMR, much like Type 4 in hummingbirds.
Hirundapus giganteus and H. caudacutus are somewhat different. In these
species the humeral tendon of TPB attaches slightly distal to the process of EMR
or on its distal edge. However, the extent of separation between that tendon
and the aponeurosis of origin of EMR is not nearly as great as it is in Types A
and B.

The use of Hirundapus giganteus to illustrate the TPB of swifts by Lowe (1939)
was unfortunate because that species is atypical of either subfamily. Although
used to support the presence of a humeral tendon of TPB in swifts, it probably
represents a stage in the loss of that tendon.

Discussion

Separate humeral tendon of TPB.—The contention that passerine birds differ
from most other birds in having a humeral tendon of TPB that is separate from
rather than fused with the aponeurosis of origin of EMR (Fig. la, b) originated
with Garrod (1876). Lowe (1939) used this feature to ally swifts and hummingbirds
with the Passeriformes (which he broadened to include also the Capitonidae,
Indicatoridae, and Picidae). Not all passeriforms (sensu strictu) have a separate
tendon; we found it fused in Eurylaimidae and Rhinocryptidae. Garrod (1876)
said it was fused in Menuridae and Atrichornithidae but Raikow (pers. comm.)
found it separate in Atrichornis clamosus. In the Caprimulgidae we found the

416 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

i i a

:
> m |
>
f g

e
C d

Fig. 2. Selected humeri (left, caudal view) showing positions of the process of M. extensor meta-
carpi radialis (opposite black triangle) and of tubercle of M. tensor propatagialis pars brevis (white
triangle): a, Coccothraustes vespertinus; b, Cordeiles minor; c, Hemiprocne comata; d, Hemiprocne
longipennis; e, Nephoecetes niger; f, Hirundapus caudacutus; g, Apus apus; h, Glaucis hirsuta; 1,
Coeligena wilsoni; j, Chaetocercus jourdanii. Some sizes have been adjusted to make them roughly
comparable, but all are not drawn to same humerus length because that measure varies greatly in
relation to body size.

tendon to be fused in Caprimulgus but completely separate in Cordeiles. Thus
separate tendons, while probably derived within birds, have evidently evolved
independently in at least the Caprimulgidae and Passeriformes.

Comparison among passerines, caprimulgids, crested swifts, and swifts strong-
ly suggests that the degree of separation of the humeral attachments of TPB and
EMR results mainly from proximal displacement of EMR. The humeral attach-
ment of TPB also moves proximally (in an evolutionary sense), but it lags behind,
creating a gap between the attachments. Eventually it may reach the level of the
process of EMR. Such stages can be represented, as in Fig. 2, by: a, a passerine;
b, Cordeiles; c, Hemiprocne comata and mystacea; d, H. longipennis,; e, Cyp-
seloidinae; f, Apodinae (Hirundapus); and g, all other Apodinae. In this morpho-
logical series, separate tendons in b-e appear to be derived relative to the fused
tendons of most non-passerines, and the fused tendons of f and g derived relative
to the separate tendons of b—e. A somewhat parallel trend occurs in hummingbirds

VOLUME 95, NUMBER 2 417

although there is considerable varation among closely related species. Types 1
and 2 tend to have the process of EMR and the tubercle of TPB widely separated,
while in Type 3 they are usually closer together (Fig. 2h and i). In hummingbirds,
as in swifts, loss or fusion of the humeral tendon of TPB (Type 4) probably
represents a derived condition (Fig. 2j). These differences are caused mainly by
proximal migration of the TPB tubercle because the process of EMR is located
far proximally in all hummingbirds. (Any quantitative comparison of the proximal
shift of these processes should compare their distance from the distal end of the
humerus to some measure of body size rather than to humeral length because the
humerus itself has become relatively shorter, to differing degrees, in the evolution
of swifts and hummingbirds.)

Attachment of TPB on EMR.—The broad attachment of the belly of TPB on
that of EMR in both swifts and hummingbirds is stated to be unique to those
families and is used to support their placement in the order Apodiformes. How-
ever, Other birds show various approaches to the condition in swifts and hum-
mingbirds. Several passerines have only a short tendon between the bellies of
TPB and EMR (swallows, pers. obs.; many Lonchurae [Bentz 1979]). In Colius
the fleshy belly of TPB extends distally as far as EMR on the dorsal surfaces of
two tendons, and in some pigeons the belly of TPB attaches on a broad tendon
almost to EMR. Thus we see that an approach toward attachment of the belly of
_ TPB on EMR has been made in at least three orders apart from swifts and hum-
mingbirds.

Within the Apodidae a broad fleshy attachment is characteristic of the Apod-
inae, as described by Lowe (1939), who dissected only members of that subfamily.
The Cypseloidinae, however, have a narrow and tendinous attachment of TPB
(noted by us in Cypseloides, Nephoecetes, and Streptoprocne) and in the Hem-
iprocnidae there is a short tendon. Thus, if we assume that the presence of one
or more tendons between the bellies of TPB and EMR as found in most birds
was the ancestral condition for swifts, we have a morphological series within the
Apodiformes from primitive (tendon) to intermediate (tapered semitendinous bel-
ly) to advanced (broad attachment of the belly).

All hummingbirds display a broad attachment resembling that of the most spe-
cialized swifts. In the morphology of the humeral tendon, however, the more
primitive stages in hummingbirds resemble the stage seen in the less specialized
swifts. Unless the ancestral hummingbirds underwent a reversal (from stage g to
h in Fig. 2) of the evolutionary trend in the humeral tendon seen 'n swifts, fol-
lowed by another reversal to parallel the trend in swifts (Compare n-j and e-g in
Fig. 2), we conclude that the resemblance of the belly of TPB in hummingbirds
and the Apodinae is the result of either parallel or convergent evolution.

Distal tendon of TPB.—The early illustration of Patagona gigas in Garrod
(1876) and subsequent work based largely on dissections of trochiline humming-
birds of Type 3 (Beddard 1898; Cohn 1968) give the impression that the presence
of a distal tendon of TPB is found in all hummingbirds and is unique to that
family. We have seen that the ‘‘tendon’’ is a diffuse, expansive, and essentially
transparent sheet in hermits. This structure might have become stronger and
better defined (as in Type 2), and further specialized into a strong tendon (Types
3 and 4) as an expression of its increasing role in supplementing extension of the
hand by EMR. We think it unlikely that evolution proceeded in the opposite
direction because reduction of a well-defined distal tendon without a change in

418 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

its function would probably result in loss of the tendon rather than in production
of a broadly investing, diaphanous sheet. We hypothesize that Types 1, 2, and
3 represent progressive evolutionary stages in the specialization of the distal
tendon of TPB.

Systematic conclusions. —The direct connection of the belly of TPB with that
of EMR in swifts and hummingbirds proves to represent an autapomorphy in
each group. Whether these apomorphies were derived from a common ancestor
(in which a tendon was present) or arose twice in different lineages cannot be
determined from this muscle alone. Thus this aspect of the muscle gives no
definite evidence for monophyly of the Apodiformes.

The alleged passerine nature of the humeral tendon of TPB in swifts and hum-
mingbirds must be discounted because some nonpasserines have a similar form
of the tendon. Furthermore, separation of the humeral tendons of TPB and EMR
is related to a proximal shift in position of origin of EMR, which has occurred in
unrelated orders or families. Once again, this feature of TPB is not by itself a
good indicator of the ordinal affinities of either swifts or hummingbirds.

Within the Apodidae, our interpretation of the evolution of specialization of
the belly and of the humeral tendon of TPB corresponds with Brooke’s (1970)
concept of a more primitive subfamily, Cypseloidinae, and a more advanced
Apodinae. Our data are not useful in determining whether the Collocaliini, Chae-
turini, and Apodini should be regarded as tribes of a single subfamily (Brooke
1970) or as two subfamilies (Collins 1976). Unfortunately, Brooke followed Lowe
(1939) in regarding the Hemiprocnidae as more nearly passerine, and therefore
more advanced, than the Apodidae. The characters used by Lowe are, in fact,
found in many other orders besides the Passeriformes, and are thus primitive
relative to the corresponding derived states in the Apodidae. The Hemiprocnidae
should precede rather than follow the Apodidae in a linear classification.

Within the hummingbirds, our interpretation of the variation in TPB would
support a hypothesis of a primitive phaethornithine group (hermits) and a more
advanced trochiline group. Within the trochilines those genera with a dorsal distal
tendon (Type 2) are more likely to be primitive (see Appendix). The presence of
superficial muscle fibers of EMR covering part of the humeral tendon of TPB in
most hermits (Fig. 1f) might best be regarded as a specialization of an otherwise
primitive condition in hermits.

Acknowledgments

We thank David M. Niles for the loan of specimens from the Delaware Museum
of Natural History, and Edwin O. Willis for providing us with anatomical spec-
imens of Ramphodon naevius. This study was partially funded by Smithsonian
Research Award SQ3372400 to R. L. Zusi.

Literature Cited

Beddard, F. E. 1898. The structure and classification of birds.—Longmans, Green and Company,
London, 548 pp.

Bentz, G. D. 1979. The appendicular myology and phylogenetic relationships of the Ploceidae and
Estrildidae (Aves: Passeriformes).—Bulletin of Carnegie Museum of Natural History 15:1-25.

Brooke, R. K. 1970. Taxonomic and evolutionary notes on the subfamilies, tribes, genera, and
subgenera of the swifts (Aves: Apodidae).—Durban Museum Novitates 9(2): 13-24.

Buri, R. O. 1900. Zur Anatomie des Fliigels von Micropus melba und einigen anderen Coracornithes,

VOLUME 95, NUMBER 2 419

zugleich Beitrag zur Kenntnis der systematischen Steilung der Cypselidae.—Jenaische Zeit-
schrift fir Naturwissenschaft 33:361-610.

Cohn, J. 1968. The convergent flight mechanism of swifts (Apodi) and hummingbirds (Trochili)
(Aves).—Unpublished Ph.D. Dissertation, University of Michigan, Ann Arbor, Michigan.

Collins, C. T. 1976. A review of the Lower Miocene swifts (Aves: Apodidae). Jn S. L. Olson, ed.
Collected Papers in Avian Paleontology Honoring the 90th Birthday of Alexander Wetmore.—
Smithsonian Contributions to Paleobiology No. 27.

Furbringer, M. 1888. Untersuchungen zur Morphologie und Systematik der Végel, zugleich ein Bei-
trag zur Anatomie der Sttitz- und Bewegungsorgane.—2 vols.—Van Holkema, Amsterdam,
1751 pp.

Garrod, A. H. 1876. On some anatomical peculiarities which bear upon the major division of the
passerine birds, Part 1.—Proceedings of the Zoological Society of London:506—519.

George, J. C., and A. J. Berger. 1966. Avian myology.—Academic Press, New York and London,
xii + 500 pp.

Lowe, P. R. 1939. On the systematic position of the swifts (Suborder Cypseli) and hummingbirds
(Suborder Trochili), with special reference to the Order Passeriformes.—Transactions of the
Zoological Society of London 24:307-349.

Morony, J., W. Bock, and J. Farrand, Jr. 1975. Reference list of the birds of the OL. —American
Museum of Natural History, New York, 207 pp.

(RLZ) National Museum of Natural History, Smithsonian Institution, Wash-
ington, D.C. 20560; (GDB) Mount Vernon College, Washington, D.C. 20007.

Appendix

Listed below are the species dissected by us for M. tensor propatagialis pars
brevis and M. extensor metacarpi radialis. We follow the terminology of Morony
Cra 97 5).

Trochilidae

Type 1.—Ramphodon naevius, Glaucis hirsuta, Threnetes ruckeri, Phaethornis yaruqui, P. malaris,
P. ruber, P. longuemareus, Eutoxeres aquila, E. condamini.

Type 2.—Androdon aequatorialis, Doryfera johannae, D. ludovicae, Florisuga mellivora, Colibri
delphinae, C. thalassinus, C. coruscans, Anthracothorax nigricollis, A. dominicus, A. viridis, A.
mango, Eulampis jugularis, Sericotes holosericeus, Chrysolampis mosquitus, Polytmus guainumbi,
Topaza pella, T. pyra, Heliothrix barroti.

Type 3.—Campylopterus duidae, Orthorhyncus cristatus, Stephanoxis lalandi, Lophornis ornata,
L. pavonina, Popelaria sp., Chlorostilbon swainsonii, C. maugaeus, Cynanthus latirostris, Cyano-
phaia bicolor, Thalurania furcata, Panterpe insignis, Damophila julie, Lepidopyga coeruleogularis,
Hylocharis chrysura, Chrysuronia oenone, Goldmania violiceps, Trochilus polytmus, Leucochloris
albicollis, Leucippus fallax, Amazilia amabilis, A. viridigaster, A. tzacatl, Elvira cupreiceps, Chal-
ybura buffonii, Lampornis castaneoventris, Adelomyia melanogenys, Heliodoxa rubinoides, H. xan-
thogonys, H. branickii, Eugenes fulgens, Sternoclyta cyanopectus, Oreotrochilus estella, Patagona
gigas, Aglaeactis cupripennis, Lafresnaya lafresnayi, Pterophanes cyanopterus, Coeligena coeligena,
C. violifer, Ensifera ensifera, Sephanoides sephanoides, Heliangelus amethysticollis, Eriocnemus
luciani, Haplophaedia aureliae, H. lugens, Ocreatus underwoodii, Lesbia victoriae, Sappho spar-
ganura, Metallura tyrianthina, Aglaiocercus kingi, Heliomaster longirostris, Philodice evelynae, Cal-
othorax lucifer, Archilochus alexandri, Mellisuga minima, Calypte costae, Stellula calliope, Myrtis
fanny, Selasphorus rufus.

Type 4.—Calliphlox amethystina, Acestrura mulsant, Chaetocercus jourdanii.

Hemiprocnidae and Apodidae

Type A.—Hemiprocne comata.

Type B.—Cypseloidinae: Cypseloides rutilus, Nephoecetes niger, Streptoprocne zonaris, S. semi-
collaris.

Type C.—Apodinae: Collocalia brevirostris, C. whiteheadi, C. esculenta, Hirundapus caudacutus,

420 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

H. giganteus, Chaetura martinica, C. pelagica, Aeronautes montivagus, A. saxatilis, Tachornis
phoenicobia, T. squamata, Cypsiurus parvus, Apus pallidus, A. pacificus.

Other families

Columbidae: Columba nigrirostris, Columbina minuta; Cuculidae: Coccyzus erythropthalmus; Po-
dargidae: Batrachostomus septimus; Nyctibiidae: Nyctibius griseus; Caprimulgidae: Chordeiles pu-
sillus, Caprimulgus europaeus; Coliidae: Colius striatus; Trogonidae: Trogon viridis; Alcedinidae:
Chloroceryle aenea; Todidae: Todus mexicanus; Momotidae: Electron platyrhynchum; Meropidae:
Merops apiaster; Coraciidae: Coracias garrulus; Upupidae: Upupa epops; Phoeniculidae: Phoenicu-
lus purpureus; Bucerotidae: Tockus erythrorhynchus; Galbulidae: Galbula ruficauda; Bucconidae:
Chelidoptera tenebrosa; Capitonidae: Megalaima haemacephala; Indicatoridae: Indicator archipe-
lagicus; Ramphastidae: Baillonius bailloni; Picidae: Jynx torquilla, Melanerpes striatus; Eurylaimi-
dae: Smithornis capensis, Eurylaimus ochromalus, Calyptomena whiteheadi; Furnariidae: Certhiaxis
subcristata; Formicariidae: Thamnophilus punctatus, Myrmotherula hauxwelli; Rhinocryptidae: Sce-
lorchilus rubecula, Scytalopus latebricola; Cotingidae: Pachyramphus cinnamomeus; Pipridae:
Chiroxiphia linearis; Tyrannidae: Ochthoeca rufipectoralis, Tyrannus dominicensis, Todirostrum
cinereum, Sublegatus arenarum; Alaudidae: Lullula arborea; Hirundinidae: Progne dominicensis,
Riparia riparia; Laniidae: Lanius cristatus; Muscicapidae: Niltava grandis; Nectariniidae: Nectarinia
jugularis; Meliphagidae: Melidectes fuscus; Parulidae: Basileuterus coronatus; Sturnidae: Acridoth-
eres tristis; Corvidae: Garrulus glandarius.

ERRATA FOR VOLUME 94(4)

page 1229 (Table 2, line 4)
**Second maxilliped,’’ should read ‘‘Second maxilliped exopod,
page 1229 (Table 2, line 6)
‘Third maxilliped,’’ should read *‘Third maxilliped exopod,”’
page 1231 (Table 3, line 10)
‘*Second maxilliped,’’ should read ‘‘Second maxilliped exopod,”’
page 1231 (Table 3, line 12)
‘Third maxilliped,’’ should read *‘Third maxilliped exopod,”’

9°

PROC. BIOL. SOC. WASH.
95(2), 1982, p. 421

JASCOTTELLA, NOM. NOV. FOR MAMILLA
SCOTT, 1974 (MICROPROBLEMATICA)
NON FABRICIUS, 1823 (MOLLUSCA)

Richard W. Huddleston and Drew Haman

Scott (1974) erected the genus Mamilla (type-species M. hemispherica Scott,
1974) for problematic microfossils of possible foraminiferal affinity from the Has-
lam Formation, Vancouver Island.

The generic name Mamilla previously had been used twice, Mamilla Fabricius,
1823 and Mamilla Wagner, 1907, both in the Mollusca. The use of this generic
name has priority in Fabricius (1823). Baker (1954) replaced Mamilla Wagner,
1907 (non Fabricius, 1823) with Weinlandella. Mamilla Scott, however, remains
a junior homonym and requires a new name (International Code of Zoological
Nomenclature, Article 53). The new name Jascottella is proposed to replace
Mamilla Scott, non Fabricius. (Repeated attempts to contact Dr. Scott in con-
formity with ICZN Appendix A, 3 have been unsuccessful.)

__ Jascottella is constructed in honor of J. A. Scott for his recognition of this new
genus.

- Acknowledgments
We thank Chevron Oil Field Research Company for their assistance and per-
mission to publish.
Literature Cited

Baker, H. B. 1954. New subgeneric names in Helicinidae.—The Nautilus 67:139-140.

Fabricius, O. 1823. Pp. 51-114 in Fortegnelse over afg. Biskop Fabriciusses etterladte Naturalier.

Scott, J. A. B. 1974. The Foraminifera of the Haslam, Qualicum, and Trent river Formations, Van-
couver Island, British Columbia.—Bulletin of Canadian Petroleum Geology 22(2):119-176.

Chevron Oil Field Research Company, P.O. Box 446, La Habra, California
90631.

ra Sy i Pe | ae

‘ » hat y ui ; 9 (ce, 4 .* b “pA

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CONTENTS

Atrophecaecum lobacetabulare, n. sp. (Digenea: Cryptogonimidae: Acanthostominae) with

discussion of the generic status of Paracanthostomum Fischthal and Kuntz, 1965, and —

Ateuchocephala Coil and Kuntz, 1960 Daniel R. Brooks and Janine N. Caira
A partial revision of the genus Notomastus Polyendeny, Capitellidae) with a description of

a new species from the Gulf of Mexico R. Michael Ewing 2

Some species of Onuphis (Polychaeta: Onuphidae) from the Atlantic Ocean Kristian Fauchald

A new species of the genus Cambarincola (Clitellata: Branchiobdellida) from Illinois. with ©

remarks on the bursa of Cambarincola vitreus Ellis, 1919, and the status of Sathodrilus

Holt, 1968 Perry C. Holt 2

A new species of oreohelicid land snail from the San Agustin plains, New Mexico

Celinda R. Crews and Artie L. Metcalf ;

A new tree snail, genus Drymaeus (Bulimulidae) from southeastern Peru

| Fred G. Thompson and Jane E. Deisler a
Yochelsoniella, nom. nov., a new name for Ellisella Rohr, 1980 (Gastropoda) non Gray, —

1858 (Coelenterata) David M. Rohr and Richard W. Huddleston
Pycnogonida of the western Pacific islands I. The Marshall Islands _ GC: Allan cae
Three new species of Pycnogonida from Sagami Bay, Japan

Koichiro Nakamura and C. ies Child

A gynandromorph of the Japanese pycnogonid Anoplodactylus gestiens (Ortmann)

The entocytherid ostracod fauna of northern Georgia
Horton H. Hobbs, Jr. and Daniel J. Peters

Two new Bs of leptomysinid mysids (Crustacea: Mysidacea) from southern California
Linda G. Gleye

The status of Cirolana parva Hansen, 1890 (Crustacea: Isopoda: Cirolanidae) with notes on
its distribution Niel L. Bruce and Thomas E. Bowman
Description of the male and notes on the female of Argeiopsis inhacae (Crustacea: Isopoda:

Bopyridae) Daniel L. Adkison, Richard W. Heard, and Guy T. Clark —

A diagnosis of the Hobbsi group, with descriptions of Caecidotea teresae, n. sp., and C.
macropropoda Chase and Blair (Crustacea: Isopoda: Asellidae) Julian J. Lewis
Gonodactylus insularis, a new stomatopod crustacean from Enewetak Atoll, Pacific Ocean

Raymond B. Manning and Marjorie L. Reaka

Rissoides, a new genus of stomatopod crustacean from the east Atlantic and South Africa
Raymond B. Manning and Ch. Lewinsohn
New records of pinnotherid crabs from the Gulf of California (Brachyura: Pinnotheridae)
Mary K. Wicksten
Studies of Neotropical caddisflies, XXXII: the immature stages of Macronema variipenne
Flint & Bueno, with the division of Macronema by the resurrection of Macrostemum
(Trichoptera: Hydropsychidae) Oliver S. Flint, Jr. and Joaquin Bueno-Soria
Notes on distribution of some Latin American cotton-stainers (Dysdercus: Pyrrhocoridae:
Hemiptera) and remarks on the biology of Dysdercus urbahni Schmidt
Joachim Adis and Richard C. Froeschner

Two new species of the frog genus Hylodes from Caparao, Minas Gerais, Brasil (Amphibia:

Leptodactylidae) W. Ronald Heyer
A new cave Platymantis (Amphibia: Ranidae) from the Philippine Islands
Walter C. Brown and Angel C. Alcala
A new species of Sphaerodactylus (Reptilia: Sauria: Gekkonidae) from eastern Cuba
Orlando H. Garrido and Albert Schwartz
Cyrtocara liemi, a previously undescribed paedophagous cichlid fish (Teleostei: Cichlidae)
from Lake Malawi, Africa Kenneth R. McKaye and Catherine Mackenzie
Hypselognathus horridus, a new species of pipefish (Syngnathidae) from South Australia

C. E. Dawson and C. J. M. Glover |

Lateral line of Diplospinus multistriatus (Teleostei: Gempylidae) Izumi Nakamura
Variation of a muscle in hummingbirds and swifts and its systematic implications
Richard L. Zusi and Gregory Dean Bentz
Errata for Volume 94(4)
Jascottella, nom. nov. for Mamilla Scott, 1974 (Microproblematica) non Fabricius, 1823
(Mollusca) Richard W. Huddleston and Drew Haman

C. Allan Child and Koichiro Nakamura ,

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(ISSN 0006-324X)

\CIETY

5 October 1982 _ _ Number 3

THE BIOLOGICAL SOCIETY OF WASHINGTON

1981-1982
Officers

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PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 423-427

SYSTEMATIC RESOLUTION OF THE
GENERA OF THE CRINIA COMPLEX
(AMPHIBIA: ANURA: MYOBATRACHIDAE)

W. Ronald Heyer, Charles H. Daugherty, and Linda R. Maxson

Abstract.—Re-evaluation of morphological analyses and generic taxonomy of
Australian myobatrachine frogs in the Crinia complex is undertaken in light of
new genetic data. All species currently in the genera Ranidella and Crinia are
assigned to Crinia. The genera Paracrinia and Geocrinia are retained.

Four Australian genera, Ranidella, Crinia, Paracrinia, and Geocrinia (Myo-
batrachidae: Myobatrachinae), collectively designated the ‘‘Crinia complex,’’
comprise a group of small, rather nondescript frogs (Heyer & Liem 1976). The
species of this complex have been partitioned into genera by recent authors with
differing results. All recent authors agree concerning the smallest groups of species
clusters in this complex, but considerable disagreement exists regarding the as-
signment of these groups at the generic level. Currently hypothesized groups and
classifications are summarized in Table 1.

The following morphological and behavioral characters differentiate the species
clusters: vomer and vomerine teeth, omohyoideus muscle, outer metatarsal tu-
bercle, belly texture, egg placement, and mating call. However, clustering algc-
rithms provide no unequivocal pattern of relationships among these species: ““There
is nO way to group the . . . taxa so that two of the derived states of these char-
acters define the same assemblage. Rather, a grouping which results in a cluster
having all the taxa with the same derived state of one character leads to conver-
gence of states in the other characters’’ (Heyer and Liem 1976:9). Thus, the data
presented by Heyer and Liem (1976) are certainly open to alternate clustering
interpretations than theirs. The phenetic analyses of Blake (1973) and Thompson
(1981) recognize the same basic species groups, but the clustering pattern of the
groups was highly variable dependent on data scoring and the algorithm used.
Within the Crinia complex, morphological variation is so limited that it has been
impossible to achieve a stable clustering scheme and, hence, taxonomic consen-
sus. For these cases where the nature of the morphological data preclude a de-
finitive analysis of relationships, use of a different data base is required for anal-
ysis of relationships.

Daugherty and Maxson (in press) recently estimated genetic relationships among
species of the Crinia complex based on MC’F (micro-complement fixation) data
from the serum protein albumin. These genetic data, in concert with the mor-
phological data, provide a new basis for determining evolutionary lineages within
the complex. The major lineages are herein proposed as generic units in order to
provide a stable classification for this complex.

The Genetic Data

Immunological distances derived from comparisons of serum albumins provide
both cladistic information and a time framework for interpreting evolutionary

424

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Species clusters and generic assignment of the Crinia complex.

Blake, 1973 Heyer & Liem, 1976 Thompson, 1981 This study
georgiana Together with Crinia Not studied Crinia
haswelli, Cri-
nia
haswelli Together with Paracrinia Not studied Paracrinia
georgiana, Cri-
nia
laevis cluster Geocrinia Geocrinia Not studied Geocrinia
signifera cluster Species group of Ranidella Distinct species Crinia
Ranidella group of Ran-
idella
riparia Together with Together with Second distinct Crinia
tasmaniensis, tasmaniensis, species group
second species Australocrinia of Ranidella
group of Rani-
della
tasmaniensis Together with ri- Together with ri- Third distinct Crinia

paria, second
species group

paria, Austral-
ocrinia

species group
of Ranidella

of Ranidella

relationships. In the past decade, such protein data have been used extensively
in phylogenetic studies of diverse amphibian taxa (e.g., Heyer and Maxson 1982;
Maxson 1981). Albumin immunological distances (ID) have been shown to esti-
mate sequence differences in albumins between species (Maxson and Wilson
1974) and to accumulate measurable sequence differences at an approximate rate
of one substitution per lineage per million years (Wilson et al. 1977). Daugherty
and Maxson (in press) have measured a series of immunological distances among
the albumins of many members of the Crinia complex. The data consist of one-
way comparisons to signifera, currently assigned to the genus Ranidella (Table
2). The pattern of divergence from signifera reveals the major genetic lineages
within this complex. |

Members of the signifera cluster (Table 2) exhibit ID values ranging from 24
to 40. The distance to riparia is 15 units and to tasmaniensis is 53 units. Clearly,
riparia belongs to the same genetic lineage as other members of the signifera
cluster. The ID value for tasmaniensis is somewhat higher than values measured
to members of the signifera group, but not as large as values to other lineages
(haswelli and laevis; see below) within the Crinia complex. Furthermore, an ID
value of around 50 is often seen between species within other frog genera (e.g.,
Maxson and Wilson 1975; Heyer and Maxson 1981). The immunological evidence
thus supports Thompson’s (1981) proposal that Australocrinia (1.e., riparia and
tasmaniensis) be synonymized with Ranidella (i.e., the signifera cluster).

The ID to georgiana is 29 units, suggesting that georgiana is part of the same
genetic lineage as the frogs in the signifera cluster. For both georgiana and
riparia, the ID values to signifera are smaller than most ID values measured
between signifera and other members of the signifera group. The taxonomic

VOLUME 95, NUMBER 3 425

Table 2.—Albumin immunological distances between signifera and other species of the Crinia
complex.*

Species compared ID to signifera
signifera cluster:
signifera 0
glauerti 24
parinsignifera 24
bilingua 30
sp. nov. 31
remota 31
deserticola 40
riparia IS)
tasmaniensis 53
georgiana 29
victoriana 133
haswelli 140

* Data from Daugherty and Maxson (in press).

conclusions are that Ranidella (including Australocrinia) and Crinia are conge-
neric and that Ranidella is a synonym of Crinia.

On the other hand, the ID value of signifera to victoriana, the only member
of the /aevis group tested to date, is 133. This very large value is concordant with
recognition of the genus Geocrinia. The ID of signifera to haswelli is similarly
high, 140. The similar, but high, ID values of haswelli and victoriana to signifera
indicate a distant relationship of haswelli and victoriana to signifera. The values
do not indicate what the relationship of haswelli is to victoriana; it could be close
or distant. In fact, preliminary data (Maxson and Daugherty, unpublished) indi-
cate a distant relationship between those taxa (ID value between 90 and 100).
These data are consistent with recognition of the genera Geocrinia (including
victoriana) and Paracrinia (including haswelli).

Discussion

Several conclusions regarding evolution of the Crinia complex logically follow
from recognition of the genera Crinia, Geocrinia, and Paracrinia.

Several of the character states that differentiate among the species clusters
have apparently evolved independently several times. Loss of the vomer and
vomerine teeth has occurred within both Crinia and Geocrinia. Both smooth and
granular bellied frogs occur in the genus Crinia. All major variation in mating
call occurs within the genus Crinia. Perhaps most notable is that variations in
life history occur within, rather than among lineages. The change from a lotic to
lentic egg placement and larval morphology has taken place entirely within the
genus Crinia, and these life history differences cannot be used to define generic
units. A similar situation occurs within the genus Geocrinia, with evolution of
terrestrial larvae from pond larvae. .

The morphological characters that differentiate among the genetically defined
lineages are, for the most part, characters involving reduction or loss as the
derived state. Crinia species have an outer metatarsal tubercle; Geocrinia species

426 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

lack the tubercle. Crinia species have an omohyoideus muscle; Paracrinia lacks
the muscle. Geocrinia species lack an outer metatarsal tubercle and toe fringing:
Paracrinia has a metatarsal tubercle and toe fringing. Of these characters, the
only one that does not involve reduction or loss as the derived state is the meta-
tarsal tubercle (Heyer and Liem 1976).

Derived states involving reduction or loss are usually considered to contain
little or no phyletic information (Hecht and Edwards 1976). That morphological
states of loss or reduction are the primary criteria permitting morphological dis-
crimination of the genera within the Crinia complex (which the genetic data show
to be greatly differentiated) suggests that the use in systematics of characters of
loss and reduction needs re-evaluation. It also further documents the extreme
morphological conservatism so often observed in anuran evolution (Maxson and
Wilson 1975; Wilson et al., 1977) and reinforces the need to examine genetic and
other categories of characters when conducting phylogenetic studies (Blake 1973).

Generic Redefinition

Crinia is the only genus requiring redefinition. The format used is comparable
to that of Heyer and Liem (1976), which may be referred to for definitions of the
genera Geocrinia and Paracrinia.

Crinia Tschudi, 1838

Synonyms .—Ranidella Girard, 1853
Camariolus Peters, 1863
Pterophrynus Lutken, 1863
Pterophryne Gunther, 1867
Australocrinia Heyer and Liem, 1976.

Type species.—Crinia georgiana Tschudi, 1838.

Diagnosis.—A myobatrachine genus; cervical cotyles widely separated; vomer
and vomerine teeth present or absent; narrow sacral diapophyses; depressor man-
dibulae muscle with or lacking a slip from the dorsal fascia; omohyoideus muscle
present; tympanum present; belly smooth or granular; toes with or without fringe;
outer metatarsal tubercle present; eggs placed in water or on land; pond or stream
larvae.

Content.—Crinia bilingua, deserticola, georgiana, glauerti, insignifera, par-
insignifera, pseudinsignifera, remota, riparia, signifera, perhaps sloanei (see
Thompson 1981), subinsignifera, tasmaniensis, tinnula, and undescribed species.
Present knowledge of variation within this genus (as here defined) does not sup-
port the recognition of species groups (also see Daugherty and Maxson, in press).

Acknowledgments

We thank P. Baverstock and his family for assistance in collecting sufficient
C. signifera to produce an antibody. We thank the many other people who pro-
vided us with specimens. Portions of this work were carried out at the Institute
of Medical and Veterinary Science in Adelaide, Australia. We thank M. Krieg,
P. Baverstock, M. Adams, and C. Watts for stimulation and support while in
Adelaide. Financial support came in part from NSF grant INT 79-24146 to LRM
and NIH grant 1-F32FM06788-01 to CHD. i

VOLUME 95, NUMBER 3 427

Literature Cited

Blake, A. J. D. 1973. Taxonomy and relationships of Myobatrachine frogs (Leptodactylidae): A
numerical approach.—Australian Journal of Zoology 21:119-149.

Daugherty, C. H., and L. R. Maxson. [In press.] A biochemical assessment of the evolution of
myobatrachine frogs.—Herpetologica.

Hecht, M. K., and J. L. Edwards. 1976. The determination of parallel or monophyletic relationships:
the proteid salamanders—a test case.—American Naturalist 110:653—677.

Heyer, W. R., and D. S. Liem. 1976. Analysis of the intergeneric relationships of the Australian

frog family Myobatrachidae.—Smithsonian Contributions to Zoology 233:1-29.

, and L. R. Maxson. 1982. Distributions, relationships, and zoogeography of lowland frogs:

The Leptodactylus complex in South America, with special reference to Amazonia. Pages 375—

388 in G. T. Prance (ed.), Biological Diversification in the Tropics. Columbia University Press,

New York. 714 pp.

Maxson, L. R. 1981. Albumin evolution and its phylogenetic implications in African toads of the

genus Bufo.—Herpetologica 37:96—104.

, and A. C. Wilson. 1974. Convergent morphological evolution detected by studying proteins

of tree frogs in the Hyla eximia species group.—Science 185:66—68.

, and 1975. Albumin evolution and organismal evolution in tree frogs (Hylidae).—

Systematic Zoology 24:1-15.

Thompson, M. B. 1981. The systematic status of the genus Australocrinia Heyer and Liem (Anura:
Leptodactylidae).—Australian Journal of Zoology 29:93-102.

Wilson, A. C., S. S. Carlson, and T. J. White. 1977. Biochemical evolution.—Annual Review of
Biochemistry 46:573-639.

(WRH) Department of Vertebrate Zoology, Smithsonian Institution, Washing-
ton, D.C. 20560. (CHD & LRM) Department of Genetics and Development, Uni-
versity of Illinois at Urbana-Champaign, Urbana, Illinois 61801.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 428-439

MEXICAN FRESHWATER SILVERSIDES (PISCES:
ATHERINIDAE) OF THE GENUS ARCHOMENIDIA,
WITH THE DESCRIPTION OF A NEW SPECIES

Barry Chernoff and Robert Rush Miller

Abstract.—Archomenidia marvelae new species, is described from two local-
ities in the basin of the Rio Papaloapan, Oaxaca and Veracruz, Mexico. Archo-
menidia marvelae differs from A. sallei (the only other species in the genus) in
several mensural characters, head pores and teeth on second pharyngobranchial,
but most notably by its smaller eye. Distinguishing features of the genus are given,
multivariate morphometrics (allometries and shape differences) of both species
are compared, and descriptive data for A. sallei are provided.

The fish faunas of the rios Papaloapan and Coatzacoalcos are biogeographically
allied with those of Central America (Rosen 1979; Miller 1982) and have been
designated as part of the large Usumacinta Fish Province (Miller 1966, 1982;
Bussing 1976). Three related genera of silverside fishes—Archomenidia, Melan-
iris, and Xenatherina—inhabit the fresh and brackish waters of this region, some-
times sympatrically, and extend well into upland areas.

Archomenidia is confined, as far as is known, to the basins of the rios Papaloa-
pan and Coatzacoalcos, in Oaxaca and Veracruz, Mexico. These rivers flow into
the Gulf of Mexico (Fig. 1). The genus was thought to contain two species, one
restricted to each drainage, but Chernoff (1981) demonstrated that A. bolivari
Alvarez and Carranza (1952) is conspecific with A. sallei (Regan, 1903).

In this paper, we describe a new species of Archomenidia from the Rio Pa-
paloapan drainage (Fig. 1), thus raising to two the number of species in the genus
and in that basin. The new form is sympatric with A. sallei at both of its known
localities. ;

Methods

Most counts and measurements are defined in Hubbs and Lagler (1964), with
the following exceptions and additions. Median lateral scale rows were counted
as described by Barbour (1973). Predorsal scales were enumerated along the
middorsal line, including the horseshoe-shaped scale at the base of the spinous
dorsal fin and the enlarged scale at the base of the occiput. Predorsal circumfer-
ential scales represent the number of scale rows around the body just anterior to
the pelvic fin base. The number of transverse scale rows includes those rows
between the middle of the anal fin base (excluding crowded, highly modified scales
at the fin base) and the midline row between the dorsal fins, as per Miller and
Carr (1974). Total gill rakers were counted on the first arch of the right side. The
vertebra of origin of a median fin has been described and figured by Chernoff et
al. (1981), and is the centrum above or below which a median fin originates. The
number of anal rays anterior to the spinous or second dorsal fin includes the
Spine; a count of zero indicates that the spinous dorsal fin is in advance of the

VOLUME 95, NUMBER 3 429

O-A.marvelae

@ -A.sallei

96° 95° 94°

Fig. 1. Map of southern Mexico showing the sample localities for Archomenidia marvelae (circles
with open stars) and A. sallei (solid circles). The type locality for A. marvelae is the Rio Bravo just
below the barrier falls, Salto de Eyipantla, and is indicated by an arrow (R. = Rio, L. = Lago).

anal fin. Head depth is measured vertically over the eye. Anal fin origin to spinous
or second dorsal fin origin refers to diagonal measurements between these land-
marks (anal fin—D1, anal fin—D2, in tables). Postdorsal fin length is measured from
the posterior margin of the second dorsal fin base to the caudal fin base.
Terminology used for the cephalic sensory canal and pore systems is modified
slightly from that of Branson and Moore (1962). The supraorbital canal includes
the canals and external pores associated with the frontal and nasal bones, in-
cluding the pore associated with the posterior orifice of the canal (postocular
sinus of Branson and Moore 1962). The temporal canal originates in the pterotic
(termed postocular commissure by Branson and Moore 1962) and continues mem-
branously until it reaches the bony canal on the posttemporal. The temporal canal

430 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

communicates with the supraorbital and dermosphenotic canals anteriorly, with
the preoperculomandibular canal midway, and with the supratemporal canal pos-
teriorly. Because atherinids have an incomplete series of infraorbital bones (and
associated sensory canals), it is necessary to divide the infraorbital canal into
preorbital (anterior infraorbital canal) and postorbital (posterior infraorbital canal)
portions. For the genus Archomenidia, the infraorbital bones consist of the lac-
rimal and jugal anteriorly, and the dermosphenotic posteriorly.

For morphometric analyses the raw data were transformed to logarithms (base
10); character designations in the morphometric section refer to logarithms of
measurements. Analysis of covariance was used to test the null hypothesis that
regressions calculated within species are not significantly different from each
other (see Snedecor and Cochran 1980). To estimate shape differences between
taxa, principal components were calculated from the covariance matrix of log-
transformed variables. The result was that two distinct species clusters, or growth
ovals, resided obliquely in the plane formed by the first two principal components,
indicating that both components contained information about size and shape. The
method of Humphries eft al. (1981) was used to regress size from the second
component, forming a size-independent shape discriminator for which loadings
and scores can be calculated.

The following institutional codes are used: IPN—Instituto Politécnico Nacio-
nal, Mexico City, Mexico; TC WC—Texas Cooperative Wildlife Collection, Texas
A&M University; TU—Tulane University, UMMZ—Museum of Zoology, Uni-
versity of Michigan; USNM—National Museum of Natural History.

Archomenidia Jordan and Hubbs, 1919

This genus is a member of the Menidiinae Schultz (1948), and may be distin-
guished from other genera in this subfamily by the following characters (many of
which were noted by Jordan and Hubbs 1919, and Schultz 1948): precaudal ver-
tebrae usually 22 or 23, rarely 21; two widely spaced prenarial sensory pits on
top of snout; anus positioned anteriorly, between anal fin origin and pelvic fin
base; upper hypural plate divided; spinous dorsal fin slightly anterior to or just
posterior to origin of anal fin; anal fin sheath short, with single row of scales,
restricted to anterior portion of fin; gill rakers on first arch fewer than 19; axillary
scale of pelvic fin well developed; ascending process of premaxilla short and
triangular; teeth in outer row of premaxilla somewhat enlarged and conical; in-
terneural elements continuous between dorsal fins; vomer without teeth; coronoid
process of dentary distinctly elevated; peritoneum dark; and mesovarium very
darkly pigmented, appearing black in preserved specimens. The type-species is
Atherinichthys sallei Regan.

Archomenidia marvelae, new species
Figure 2

Holotype.—UMMZ 209012 (formerly TCWC 1848.5), adult female, 80.2 mm
SL, Mexico, Veracruz, Rio Bravo just below barrier falls, Salto de Eyipantla;
collected by Conner, Kent and Meyer, 19 July 1962.

Paratypes.—UMMZ 209013, 2 specimens, 45.8—94.7 mm SL, collected with
holotype. UMMZ 179892, 41 specimens (including 5 cleared and stained), 12-

VOLUME 95, NUMBER 3 431

Fig. 2. Archomenidia marvelae, holotype, 80.2 mm SL, UMMZ 209012. Photo by E. C. Theriot.

36.2 mm SL, type locality; O. P. Johnson, J. Hamilton and D. C. Robinson, 23
June 1961. UMMZ 209850, 17 specimens, 59.1-86.2 mm SL, Mexico, Oaxaca,
Rio Papaloapan at Papaloapan, ca. 100 m below Hwy 145 bridge; R. R. & F. H.
Miller and B. Chernoff, 25 January 1982.

Diagnosis.—A species of Archomenidia that differs from A. sallei in having a
smaller eye ($9.5 vs. 29.8% SL; x = 8.6 vs. 11.1% SL); a more slender body
(greatest depth, x = 19.3 vs. 22.3% SL); a shorter preanal distance (x = 57.1 vs.
60.2% SL); anal fin usually originating under 18th vertebra (vs. usually 19th); a
shorter diagonal distance between origins of anal and spinous dorsal fins (« =
' 18.9 vs. 22.0% SL); a shorter distance between origins of anal and second dorsal
fins (x = 22.4 vs. 24.7% SL); shorter pelvic fins (x = 14.1 vs. 15.5% SL); stripe
on ventral surface of caudal peduncle with two or three rows of large melano-
phores (vs. dusky stripe, large melanophores not organized into rows); relatively
few teeth on dentary, in one or occasionally two rows (vs. relatively many teeth,
crowded in two or three rows); teeth on second pharyngobranchial with shoulder
posteriorly (vs. conical teeth, lacking shoulder); and anterior infraorbital canal
with four or five pores (vs. two or three pores).

Description.—Data on body proportions appear in Table |. Meristic values for
the holotype are designated with asterisks below.

Body somewhat elongate, attaining 94.7 mm SL, and neither very deep-bodied
anteriorly nor extremely compressed laterally. Head with blunt, broad, rounded
snout; smooth, slightly convex interorbital region, and thickened lips. Anal and
second dorsal fins strongly falcate. Pectoral fin inserted below lateral stripe, short,
not reaching beyond vertical from mid-point of pelvic fins. Lower lobe of caudal
fin longer and broader than upper lobe.

Scales lacking circuli or radii on exposed posterior field, with only three to four
radii on anterior field, and moderately imbricate on sides of body. Posterior scale
margins of adults entire to crenate on sides of body, and crenate or with one or
two teeth on predorsal scales; juveniles with entire to slightly crenate scales.
Scales present on base of caudal fin, extending distally along membrane between
outer rays; procurrent caudal rays covered by scaly sheath, becoming thickened
in adults.

First dorsal fin spines 4*(25 counts), 5(23), 6(1). Second dorsal fin rays I1,8(9),
[,9*(33), I,10(7). Anal fin rays I,16(1), I,17(13), I,18*(27), I,19(7), 1,20(1). Anal fin
elements anterior to: second dorsal fin origin 9(10), 10*(22), 11(14), 12(3); spinous
dorsal fin origin 0(33), 1(2), 2*(2), 3(1). Pectoral fin rays 13*(3), 14(19), 15(25),
16(2). Lateral scale rows 37(5), 38*(8), 39(28), 40(6), 41(2). Predorsal scales 17(7),

432 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1. Morphometric data of Archomenidia marvelae (n = 33-49) and A. sallei (n = 51-77).
Values are in thousandths of SL.

A. marvelae A. sallei
Holotype Range x Range x

SL, mm 80.2 24.5—94.7 47.1 26.8-75.7 51.4
D1 to snout tip 581 543-596 567 564-604 584
D2 to snout tip 718 664-718 698 689-739 716
Preanal length 588 544-597 571 574-636 602
Prepelvic length 382 379-421 399 400-438 418
Prepectoral length 231 231-261 253 253-287 270
Head length 226 226-269 250 250-290 267
Head depth 113 112-139 122 125-145 134
Eye diameter 71 71-95 86 98-132 111
Snout length 65 51-76 64 60-77 69
Interorbital width 77 73-92 81 77-95 86
Body depth 208 168-218 193 200-234 223
Body width 107 65-107 81 68-111 86
Caudal-peduncle

Length 184 173-208 192 171-216 187

Width 47 21-51 28 27-51 37

Least depth 89 73-92 84 87-103 95
Postdorsal length 188 187-215 203 180-225 200
Anal fin—D1 209 158-216 189 198-234 220
Anal fin—D2 248 197-254 224 229-266 247
Anal fin base 258 220-268 244 224-258 240
D2 fin base 106 83-116 102 95-122 107
Pelvic fin length 146 121-157 141 143-166 155
Pectoral fin length 222. 197-245 Oe, 221-259 239

18(21), 19*(14), 20(5), 22(2). Predorsal circumferential scales 20(1), 21(8), 22*(39),
23(1). Transverse scale rows 7*(46), 8(3). Scales around caudal peduncle 12*(49).
Total gill rakers 13(1), 14(6), 15(18), 16*(10), 17(4). Vertebrae: total 39(2), 40*(35),
41(2); precaudal 21(1), 22(21), 23*(9); and caudal 16(1), 17*(9), 18(26), 19(3). Ver-
tebra of origin of: spinous dorsal fin 17(2), 18(32), 19*(6); second dorsal fin 24(5),
25*(35), 26(1); and anal fin 17*(3), 18(24), 19(11).

Cephalic sensory system: supraorbital canal with five pores; temporal canal
with two pores; supratemporal canal with one pore; preoperculomandibular canal
with 14 pores; anterior infraorbital canal with four or five pores; and posterior
infraorbital canal with one pore.

Pigmentation.—Overall appearance of adults dusky to dark above and lighter
below; smaller specimens generally lighter. Superficial melanophores peppered
over top of head, not very concentrated in posterior two-thirds of head. Brain
pigment heart-shaped, dark, and with moderately incised notch. Melanophores
becoming larger and more concentrated over snout. Premaxilla darkly pigmented
with small melanophores. Dentary, gular, and preorbital regions pigmented, with
melanophores ending before anterior rim of orbit. Preopercle, subopercle, bran-
chiostegal membranes, and remainder of intermandibular region generally im-
maculate. Upper half of opercle with patch of melanophores.

Dorsum dusky, with prominent pre-, inter- and postdorsal stripes. Predorsal

VOLUME 95, NUMBER 3 433

86 @ A
t )
@-A.sallei @) @ at
e%e A
A-A.marvelae At

log Eye Diameter

1.4 jegusu 2.0

Fig. 3. Plot of log eye diameter against log SL for Archomenidia marvelae (n = 49) and A. sallei
(n = 77). Each symbol represents at least one individual.

stripe with large melanophores loosely organized into three rows. Scales above
lateral stripe with pigment concentrated along scale margins giving cross-hatched
appearance; scales along dorsal fin bases entirely covered with melanophores.
Lateral stripe uniform, its width less than height of scale, originating at pectoral
insertion, expanding very little at caudal base. Pigment reduced on scales below
lateral stripe; these scales faintly outlined for several rows anteriorly, grading to
one row on caudal peduncle. Pigment reduced or absent on lower flanks, breast
and belly. Some individuals with flecks of pigment between tips of pelvic fins and
base of anal fin. Row of melanophores along base of anal fin, beginning at mid-
point of fin and continuing on ventral side of caudal peduncle up to procurrent
caudal rays; ventral caudal peduncle stripe with two or three pigment rows.

Melanophores present along spines of anterior dorsal fin; interradial membranes
clear. Anterior lobe of second dorsal fin blackened with large flecks of pigment
on rays and membranes of first five or six elements, beyond which pigment re-
stricted to distal portion. Anal fin with prominent black anterior lobe, large me-
lanophores on and between first six elements; remainder of fin peppered along
distal margin. Outer rays of caudal fin dusky to dark; small melanophores forming
black marginal band; base of caudal fin dusky, grading into lighter central region.
Pectoral fin with large melanophores along upper rays and base of fin. Pelvic fins
immaculate.

434 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

A.sallei

A. marvelae

Fig. 4. Scatter diagram of scores on size-independent shape factor (H) against first principal
component (I) for Archomenidia marvelae (n = 39) and A. sallei (n = 52). Each symbol represents
at least one individual.

Morphometrics.—Archomenidia marvelae and A. sallei can be readily distin-
guished with morphometric characters; descriptive ratios are given in Table 1.
The disparity between these forms results from change in proportion and allom-
etry. Eye diameter is one of the most obvious distinguishing features (Fig. 3) and
the slopes of the within-species regressions on SL are not significantly different
(P > .1). Thus, for the range of lengths examined, eye diameter scales! equiva-
lently with length, but A. marvelae has a smaller eye for any given SL (means
and intercepts are significantly different, P < .001).

Overall morphometric divergence of these taxa is evident from the plot of
scores on the first principal component and shape discriminator (Fig. 4); character
coefficients are given in Table 2. Scores on the shape discriminator are completely
nonoverlapping, and reflect the relatively smaller eye, more slender body, nar-
rower anal fin base and longer caudal peduncle of A. marvelae. Scores on this
factor are size-independent and even the smallest specimens included (24.5 mm
SL) can be discriminated.

! The verb, to scale, indicates covariation of variables and does not refer to structures of dermal
origin.

VOLUME 95, NUMBER 3 435

Table 2. Principal components analyses of Archomenidia marvelae (n = 39) and A. sallei (n = 52):
A) between species analysis with coefficients for the first two principal components and the size-
independent shape factor (PCI, PCII, and H,! respectively); B) coefficients of first principal compo-
nent within each species.

Between species Within species
Variables PCI PCII H marvelae sallei
SL .96 1.02
D1 to snout tip 2D = 15 —.14 .98 1.07
D2 to snout tip 22 == 15) —.14 .99 1.06
Preanal length se) = L7/ —.07 .99 1.09
Prepelvic length aD) —.12 eal .99 1.04
Prepectoral length a) .03 .03 93 aT
Head length 21 05 .06 .90 .99
Head depth Al 19 20 .89 92
Eye diameter 23 57 58 95 .99
Snout length 5) —103 — .02 1.10 1.07
Interorbital width .20 .07 .08 .88 .82
Body depth 26 wD aD 1.10 1.03
Caudal-peduncle
Length 21 — .42 — .42 297) 1.00
Least depth 5 lS 16 1.07 .93
~ Postdorsal length wp =. 3h) = 37) iy ES)
Anal fin—D1 Ly 23 28) 1.14 1.06
Anal fin—D2 .26 .04 .04 el 1.05
Anal fin base 22 = 33) = 32 1.01 95
D2 base length 22 —.10 = 1I(0) SH 91
Pelvic fin length 24 .07 .08 1.03 1.01

1a = —.02, B, = 1.00, B, = —.02.

Differences are also apparent in the relative covariances of variables with a
general measure of size. Coefficients for the multivariate allometry of a taxon are
calculated from the first principal component (=general size; Jolicoeur 1963) and
normalizing the coefficients to a mean-square of one (i.e., by multiplying each
loading by the square root of the number of variables). The standardized coeffi-
cients reflect the relative scaling of all variables with general size, and thus may
be used to describe a taxon (Humphries 1981). Coefficients about 1.00 scale
isometrically with size, those greater and less than 1.00 represent positive and
negative allometries, respectively. Comparison of the loadings for A. marvelae
and A. sallei (Table 2) indicates that the patterns of relative growth within each
Species are disparate. For example, head length and eye diameter are negatively
allometric in A. marvelae, whereas these characters scale isometrically in A.
sallei. To a degree, deviations of the coefficients between species account for the
shape differences noted above. The discordance in multivariate allometries can-
not be attributed to environmental effects, because two populations of A. sallei
sympatric with A. marvelae are included in the analyses, and they do not differ
significantly from the allopatric populations of A. sallei.

Two other aspects of the allometries are worthy of mention. Firstly, we noted
above that eye diameter scales equivalently with SL for both taxa (Fig. 3), but
the coefficients for these variables are not equivalent (Table 2). These observa-

436 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tions are not contradictory because relative changes in both variables to size
within each species are similar. That is, eye diameter and SL are negatively
allometric in A. marvelae, and both are isometric in A. sallei. Secondly, the
discrepancy between the relative scalings of SL with size, and the departure from
isometry for SL, point to the fallacy of using any particular length as an indicator
of size. Humphries et al. (1981) have defined general size as a factor which leaves
the smallest mean squared residual when predicting all other distance measures
within a population. For A. marvelae, SL does not predict the other variables
very well, but most importantly, during growth this species increases in overall
size more quickly than in SL.

Distribution.—Archomenidia marvelae is known from only two localities in the
basin of the Rio Papaloapan (Fig. 1). The type locality is the Rio Bravo just below
the barrier falls, Salto de Eyipantla. The Rio Bravo drains Lake Catemaco and
interestingly, no atherinids have been collected above the falls or in the lake
(Miller 1975). The other locality is in the Rio Papaloapan, upstream from the
affluence of the Rio Tonto, at the town of Papaloapan.

Habitat and Associates.—Both habitats from which A. marvelae has been
collected have strong currents and clear water, although the Rio Papaloapan may
become muddied during the rainy season. At the type locality, the stream Is ca.
25 m wide with coarse sand, rocks and boulders. At Papaloapan, the river is ca.
500 m wide with sand, fine mud, gravel bars and occasional large rocks.

These locations, part of the tropical lowlands of Oaxaca and Veracruz, are
surrounded by perennial forest and fall within the 10°C minimum and 40°C max-
imum isotherms (Rzedowski 1978); the region receives between 160 and 320 cm
of rainfall yearly.

Other fish species known from or near the type locality, below the falls, are:
Xiphophorus helleri, Archomenidia sallei, Cichlasoma ellioti, C. fenestratum,
and C. octofasciatum. The associated fish species captured with the paratypes
in the Rio Papaloapan are: Dorosoma anale, Astyanax fasciatus, Cathorops
aguadulce, Poecilia mexicana, P. sphenops, Strongylura hubbsi, Archomenidia
sallei, Agonostomus monticola, and Cichlasoma ellioti.

Etymology.—We are pleased to name this species for Marvel B. Parrington,
whose dedication and hard work have contributed significantly to ichthyological
efforts at The University of Michigan for the past 17 years. The name marvelae
is a feminine noun in the genitive singular.

Archomenidia sallei (Regan)

Archomenidia sallei has remained a poorly known taxon, receiving scant sys-
tematic attention since its description by Regan (1903). Because Chernoff (1981)
provided data relevant only to the status of A. bolivari, we present a brief de-
scription of A. sallei below, thus completing the meristic and morphometric data
base for this genus.

Mensural data appear in Table 1, and A. sallei is contrasted with A. marvelae
above. The observations below result from examination of specimens throughout
the known range of the species (Fig. 1).

Archomenidia sallei is deep bodied anteriorly, laterally compressed, attaining
75 mm SL. Head deep and broad, with conspicuously large eye (see Chernoff

VOLUME 95, NUMBER 3 437

1981, Fig. 1) and parabola-shaped snout. Anal and second dorsal fins falcate;
pectoral fin short and deep, upper rays reaching to vertical from mid-point of
pelvic fins, not greatly longer than middle pectoral rays. Teeth in both jaws
conical, curved and crowded into two or three irregular rows. In adults, exposed
margin of scales entire to crenate (occasionally with one or two laciniations in
predorsal series); posterior field of scales without circuli or radii.

First dorsal fin spines 3(1), 4(24), 5(24), 6(4). Second dorsal fin rays I,8(8),
1,9(60), 1,10(12). Anal fin rays I,15(1), I,16(3), I,17(17), 1,18(32), 1,19(22), 1,20(4).
Anal fin elements anterior to: second dorsal fin origin 8(2), 9(12), 10(28), 11(10);
spinous dorsal fin origin 0(57), 2(1), 3(1). Pectoral fin rays 13(1), 14(22), 15(23),
16(6). Lateral scale rows 36(1), 37(15), 38(25), 39(29), 40(10). Predorsal scales
15(2), 16(2), 17(23), 18(19), 19(5), 20(1). Predorsal circumferential scales 18(1),
19(2), 20(20), 21(13), 22(33). Transverse scale rows 7(52). Scales around caudal
peduncle 12(80). Total gill rakers 14(1), 15(16), 16(22), 17(12), 18(1). Vertebrae:
total 39(4), 40(40), 41(3); precaudal 22(26), 23(20), 24(1); caudal 17(23), 18(23),
19(1). Vertebrae of origin of: spinous dorsal fin 17(5), 18(36), 19(15); second dorsal
fin 24(11), 25(33), 26(3); anal fin 18(13), 19(30), 20(3).

Cephalic sensory system: supraorbital canal with five pores; temporal canal
with one or two pores (the anterior pore on the pterotic may be absent); supraor-
bital canal with one pore; preoperculomandibular canal with 14 pores; anterior
‘infraorbital canal with two or three pores; and posterior infraorbital canal with
one pore.

Material Examined.—Rio Papaloapan Drainage—IPN 249 (21 specimens: 13.5—
32.3 mm SL), stream below Salto de Eyipantla, downstream from Comapan,
Veracruz, 25 May 1951. UMMZ 92121 (18:16.8-48.7), USNM 123208 (3:36.7-
45.7), Rio Hueyapan, downstream from Hueyapan, ca. 9.7 km E Cuatotolapan,
NW Acayucan, Veracruz, 23 July 1910. UMMZ 207708 (51:40-65), Rio Hueyapan
at E end of Hueyapan, 3.1 km from Highway 180, Veracruz, 24 Jan. 1979. UMMZ
209851 (3:52.1—75.7), Rio Papaloapan at Papaloapan, ca. 100 m below Highway
145 bridge, Oaxaca, 25 Jan. 1982.

Rio Coatzacoalcos Drainage.—TU 39125 (1:68.8), Rio Jaltepec at Jesus Car-
ranza, Veracruz, 27 Jan. 1967. UMMZ 184767 (23:47.4-63.0), Rio Jaltepec, above
bridge crossing on Trans-Isthmian Highway, 72.4 km S Acayucan, Veracruz, 25
Feb. 1959. USNM 162503 (20:18.4—36.8; paratypes of A. bolivari), Rio Coatza-
coalcos en el Carrizal, Santa Maria Chimilapa, Oaxaca, 20 May 1950. UMMZ
163238 (7:26.8—29.1; paratypes of A. bolivari), Rio Grande or Rio Almoloya, El
Ocotal, Matias Romero, Oaxaca, 29 April 1950. UMMZ 178528 (12:34.6—63.7),
Rio Almoloya, 30.3 km N southern terminus of Highway 185, Oaxaca, 27 March
1957.

Resumen

Se describe Archomenidia marvelae, nueva especie de la familia Aterinidae.
Su distribucion se limita a aguas abajo del Salto de Eyipantla y en el Rio Papa-
loapan, Veracruz y Oaxaca, Mexico. La otra especie en este género, A. sallei,
se distribuye en las cuencas de los rios Coatzacoalcos y Papaloapan de los estados
de Oaxaca y Veracruz. Archomenidia marvelae se distingue basicamente de A.
sallei por un ojo mas pequeno. Otros caracteres diagnosticos son: la altura del

438 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

cuerpo, la posicion de las aletas anal y dorsales, la longitud de la aleta pélvica,
el numero de dientes mandibulares, el numero de poros de linea lateral cefalica,
asi como la pigmentacion. Estas especies tienen crecimientos alométricos muy
diferentes. Archomenidia bolivari es considerado en sinonimia con A. sallei. Los
caracteres en que distinguen a Archomenidia de otros géneros dentro de la
subfamilia Menidiinae Schultz (1948) son presentados.

Acknowledgments

We extend our appreciation to: Clyde D. Barbour (Wright State University),
Julian M. Humphries (UMMZ) and Richard E. Strauss (UMMZ) for commenting
on the mansucript; José Alvarez del Villar (IPN) for a gift of paratypes of A.
bolivari; John D. McEachran (TCWC) for a gift of holotype and paratypes of A.
marvelae; curators and staff of USNM for use of facilities, especially Susan
Jewett for her help during our visit; Edward C. Theriot (U. Michigan) for pro-
duction of Fig. 2; Victor Torres (U. Michigan) for assistance with the Spanish
summary; Robin Z. Chernoff for assistance with Fig. 1; and Frances H. Miller
for typing the manuscript and help in the field. Permission to collect fishes in
Mexico was generously granted by the Direcciodn General de Pesqueras (Permit
No. 3616). Field work was supported by the National Science Foundation
(GB-6272x, GB-14871, and DEB 77-17315, to RRM). Support from an Edwin C.
Hinsdale Scholarship, UMMZ, to BC is gratefully acknowledged.

Literature Cited

Alvarez, J., and J. Carranza. 1952. Cuatro especies nuevas de peces dulceacuicolas del sureste de
México.—Ciencia 11(10—12):281-289.

Barbour, C. D. 1973. The systematics and evolution of the genus Chirostoma Swainson (Pisces,
Atherinidae).—Tulane Studies in Zoology and Botany 18:97-141.

Branson, B. A., and G. A. Moore. 1962. The lateralis components of the acoustico-lateralis system
in the sunfish family Centrarchidae.—Copeia 1962(1):1—108.

Bussing, W. A. 1976. Geographic distribution of the San Juan ichthyofauna of Central America
with remarks on its origin and ecology, pp. 157—175.—Jn Investigations of the Ichthyofauna
of Nicaraguan Lakes, T. B. Thorson, ed. School Life Sciences, University of Nebraska, Lin-
coln, Nebraska.

Chernoff, B. 1981. Taxonomy of the Mexican atherinid fish genus Archomenidia.—Copeia 1981(4):

913-914.

, J. V. Conner, and C. F. Bryan. 1981. Systematics of the Menidia beryllina complex (Pisces:

Atherinidae) from the Gulf of Mexico and its tributaries —Copeia 1981(2):319-336.

Hubbs, C. L., and K. F. Lagler. 1964. Fishes of the Great Lakes Region.—University of Michigan
Press, Ann Arbor, Michigan, 213 pp.

Humphries, J. M. 1981. The evolution of a species flock in the genus Cyprinodon (Pisces: Cyprin-

odontidae).—Unpublished Ph.D. Dissertation, University of Michigan, Ann Arbor. 233 pp.

, F. L. Bookstein, B. Chernoff, G. R. Smith, R. L. Elder, and S. G. Poss. 1981. Multivariate

discrimination by shape in relation to size.—Systematic Zoology 30(3):291—308.

Jolicoeur, P. 1963. The multivariate generalization of the allometry equation.—Biometrics 19:497—
499.

Jordan, D. S., and C. L. Hubbs. 1919. Studies in ichthyology. A monographic review of the family
of Atherinidae or silversides.—Leland Stanford Jr. University Publications, University Series
40: 1-87.

Miller, R. R. 1966. Geographical distribution of Central American freshwater fishes.—Copeia 1966(4):

773-802.

1975. Five new species of Mexican poeciliid fishes of the genera Poecilia, Gambusia, and

VOLUME 95, NUMBER 3 439

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44.

—. 1982. Pisces—Jn Aquatic Biota of Mexico, Central America and the West Indies, S. H.

Hurlbert and A. Villalobos, eds. San Diego State University, San Diego, California, in press.

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Regan, C. T. 1903. Ona collection of fishes made by Dr. Goeldi at Rio Janeiro.—Proceedings of
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Rosen, D. E. 1979. Fishes from the uplands and intermontane basins of Guatemala: Revisionary stud-
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Rzedowski, J. 1978. Vegetacion de México.—Editorial Limusa, S.A., México, D. F., México. 432
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Schultz, L. P. 1948. A revision of six subfamilies of atherine fishes, with descriptions of new genera
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School of Natural Resources and Museum of Zoology, The University of Mich-
igan, Ann Arbor, Michigan 48109.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 440-450

A NEW SPECIES OF SKATE FROM THE WESTERN
INDIAN OCEAN, WITH COMMENTS ON THE
STATUS OF RAJA (OKAMEJET)
(ELASMOBRANCHII: RAJIFORMES)

John D. McEachran and Janice D. Fechhelm

Abstract.—Raja (Okamejei) heemstrai n. sp. is described from six specimens
captured off Kenya and Tanzania. Within Okamejei the new species most closely
resembles R. powelli in possessing a rhomboid-shaped disc, relatively long point-
ed snout and large interspace between the dorsal fins. The two species can be
distinguished from each other by several proportional measurements, number of
tooth rows in the upper jaw, coloration, and structure of neurocranium and scap-
ulocoracoids. The third Okamejei species from the Indian Ocean, R. philipi,
appears to be synonymous with R. powelli but the holotype and only known
specimen of R. philipi has been lost and its status remains uncertain. Raja (Oka-
mejei) is compared with R. (Dipturus) and it is concluded that only the structure
of the anterior margin of the anterior neurocranial fontanelle distinguishes the
two subgenera. The second known specimen of Cruriraja andamanica is briefly
described and compared with the holotype of the species.

Stehmann (1976) revised the poorly known skate fauna of the tropical Indo-
West Pacific and concluded that only 10 species occur in this vast area. He stated
that the skate genera-subgenera Breviraja (3 spp.), Cruriraja (1 sp.), Raja (Am-
blyraja) (1 sp.), R. (Dipturus) (1 sp.), R. (Okamejei) (2 spp.), R. (Rajella) (1 sp.)
and R. (Rostroraja) (1 sp.) occur in the area and that this area has served as a
migration route or passage between the Pacific and Atlantic oceans rather than
an area of colonization. Recently McEachran and Fechhelm (1982) described a
new species of Pavoraja from the eastern Indian Ocean, and herein we describe
another species of R. (Okamejei) from the western Indian Ocean. This suggests
that more intensive collecting may reveal that the deeper waters of the tropical
Indo-West Pacific support a rather diverse skate fauna. We also discuss the status
of the other two Indian Ocean R. (Okamejei) species, distinctions between R.
(Okamejei) and R. (Dipturus), and comment on the second known specimen of
Cruriraja andamanica.

Specimens of the new species were obtained from the Rhodes University,
Grahamstown, South Africa (RUSI) and the Zoologisch Museum Universiteit van
Amsterdam (ZMA). Specimens of Raja (Okamejei) powelli and Cruriraja anda-
manica were obtained from the British Museum (Natural History) (BMNH) and
ZMA respectively. One specimen each of the new species and R. powelli was
dissected to reveal the structure of the neurocranium and scapulocoracoids. The
remaining specimens were radiographed to verify the anatomical observations
based on dissections and to count vertebrae, pectoral radials and pelvic radials.
Methods for making measurements and counts follow McEachran and Compagno
(1979; 1982). 3

VOLUME 95, NUMBER 3 44]

Fig. 1. Raja heemstrai RUSI 13812 (Holotype). a. Dorsal view; b. Distolateral section of right
pectoral fin showing color pattern; c. Ventral view.

Raja (Okamejei) heemstrai, new species
(Bigs al 2)

Holotype.—RUSI 13812, 404 mm TL, adolescent male, collected off Kenya,
02°59'S, 40°25'E, 260 m, 12 December 1980, aboard R/V ‘‘Fridtjof Nasen’’ by
Philip C. Heemstra.

Paratypes.—RUSI 15920, 515 mm TL, adolescent female, 458 mm TL, 365 mm
TL juvenile females; USNM 231866, 419 mm TL, adolescent male; all collected
with holotype. ZMA 113.399, 219 mm TL, juvenile male, collected off Tanzania,
near Dar es Salaam, ca. 07°S, 40°E, 274—457 m, 26—28 June 1974.

Diagnosis.—Disc rhomboid-shaped; snout about one-half head length; preor-
bital length 14.9-17.1% of total length; orbital diameter greater than interorbital
distance; spiracle length about one-half orbit diameter; distance between first gill
slits 11.6—13.2% of total length; distance between fifth gill slits 6.9-7.9% of total
length; teeth in upper jaw 31-35; distance between dorsal fins greater than length
of first dorsal fin base; dorsal surface with numerous symmetrically arranged
ocelli, consisting of narrow yellow rings surrounding brown or tan centers.

Description.—Disc 1.2 times as broad as long (Table 1); maximum angle in
front of spiracles 80° in holotype (71°-90° in paratypes); margin of disc concave

442 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Raja heemstrai RUSI 15920 (Paratype, 515 mm TL, female), dorsal view.

from tip of snout to midlength of snout, slightly convex from midlength of snout
to level of orbits, slightly concave from level of orbits to outer corners, outer
comers narrowly rounded, posterior margins slightly concave laterally, but slight-
ly convex medially. Axis of greatest width 76% (69-75%) of distance from tip of
snout to axil of pectoral fins. Pelvic fins moderately incised, anterior lobe narrow
and acutely pointed, anterior margin 78% (71-85%) as long as distance from origin
of anterior lobe to posterior extreme of fin. Tail slender, little depressed, its width
at midlength about two-thirds diameter of eye. Tail with narrow lateral fold along
ventrolateral surface, running from axil of pelvic fin to near tip of tail. Length of
tail from center of cloaca to tip 1.2 (1.1—1.3) times distance from tip of snout to
center of cloaca.

443

VOLUME 95, NUMBER 3

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VOLUME 95, NUMBER 3 445

Table 2. Neurocranial and scapulocoracoid measurements of R. heemstrai and R. powelli ex-
pressed as percentage of nasobasal length or scapulocoracoid length.

R. heemstrai R. powelli
USNM 231866 BMNH 1939.5.24:17
419 mm TL 347 mm TL
Nasobasal length (mm) 37.8 33.3
Cranial length 245 243
Rostral cartilage length 146 140
Prefontanelle length 123 120
Cranial width 111 12
Interorbital width 35 45
Rostral base 28 25
Anterior fontanelle length 39 44
Anterior fontanelle width 15 19
Posterior fontanelle length 40 38
Posterior fontanelle width 10 17
Rostral appendix length 58 27
Rostral appendix width 20 DS
Cranial height m8 708}
Width across otic capsules 52 51
Least width of basal plate 31 35
Greatest width of nasal capsule 41 47
Internasal width 26 26
_ Scapulocoracoid length (mm) 26.3 DDS)
Scapulocoracoid height 61 56
Premesocondyle 46 38
Postmesocondyle 54 62
Postdorsal fenestra length 27) 34
Postdorsal fenestra height 19 Ili
Anterior fenestra length ap) 20
Anterior fenestra height 25 24
Rear corner 46 42

Preorbital length 4.2 (4.0-4.6) times as long as orbit; preoral length 2.5 (2.3-
2.7) times internarial distance. Interorbital distance 0.9 (0.9-1.0) times length of
orbit, orbit length 2.1 (2.0-3.0) times as long as spiracles. Anterior nasal flap
(nasal curtain) fringed along distal margin, posterior nasal flap poorly developed
and very weakly fringed. Upper and lower jaws slightly arched near symphysis.
Teeth with pointed cusps near symphysis but with rounded cusps near margin of
jaws Guvenile specimens with rounded cusps), teeth in quincunx arrangement.

Distance between first gill slits 1.8 (1.7-1.9) times as great as between nares;
distance between fifth gill slits 1.0 (1.0-1.2) times as great as between nares;
length of first gill slits 1.7 (1.4-2.2) times length of fifth gill slits and 0.2 times
mouth width. First dorsal fin about equal in size and shape to second, second
dorsal fin separated from epichordal caudal-fin lobe by distance equal to about
one-half base of second dorsal fin; epichordal lobe of caudal fin relatively well
developed, low, but base length equal to that of second dorsal fin.

Upper surface of disc, pelvics and tail devoid of dermal denticles. Ventral
surface naked. Orbit with 3 (1-4) thorns on anteromedial margin and 2 (2-4) thorns
on posteromedial margin; | thorn medial to each spiracle; 1 (2) nuchal thorn;
tail with 3 (3—5) irregular rows of thorns on dorsal surface; 2 interdorsal thorns.

446 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

icm

Fig. 4. Lateral view of scapulocoracoid. a. R. heemstrai USNM 231866 (Paratype), b. R. powelli
BMNH 1939.5.24:17, 347 mm TL, male. af—anterior fontanelle, msc—mesocondyle, mtc—metacon-
dyle, pdf—postdorsal fenestra, prc—procondyle, pvf—postventral fenestra or foramina, rc—rear cor-
ner, scp—scapular process.

Neurocranium with a relatively long and moderately slender rostral shaft (Fig.
3a, Table 2); rostral appendices flat, rather long and attached over the entire
length to rostral shaft; propterygia of pectoral girdle extending to midlength of
rostral shaft; nasal capsules small, rhomboid-shaped and set about 15° angle to
transverse axis of neurocranium; foramen for profundus nerve on leading edge
of nasal capsule; interorbital region moderately narrow (Table 2); preorbital pro-
cesses poorly developed, continuous with incised supraorbital crests, postorbital
processes poorly developed; anterior fontanelle elliptical; posterior fontanelle
narrow with slightly convex lateral margins; foramen for anterior cerebral vein
on a vertical with dorsal margin of optic nerve foramen and on a line connecting
foramina for preorbital and orbitonasal canals (Fig. 3b); foramen for trochlear
nerve dorsal to optic nerve foramen; jugal arches very slender (Fig. 3c); basal
plate and nasal plate moderately broad (Fig. 3d).

Scapulocoracoids rather low and anteroposteriorly expanded both between
procondyle and mesocondyle and between mesocondyle and metacondyle (Fig.
4a, Table 2); anterior, postdorsal and postventral fenestrae expanded; rear corner
elevated.

Claspers of holotype and adolescent male paratype not fully formed but com-
ponents appear to be present. Inner dorsal lobe possesses slit; inner ventral lobe
possesses rhipidion, shield, funnel, sentinel and spike (Fig. 5).

Color.—After preservation in formalin and storage in alcohol, dorsal surface
dark brown with numerous, fairly symmetrically arranged ocelli; ocelli bi-colored
with a narrow yellowish-beige ring surrounding a brown or tan center, or ocelli
lacking ring and appearing as a brown or tan spot. Ocelli ranging in size from
to 1 times diameter of orbit. Number of ocelli variable but arrangement similar
on all specimens examined. Smaller specimens with a more distinct dorsal pat-
tern, becoming muted in larger specimens. Margin of disc and sides of snout light
tan. Dorsal and caudal fins dark brown. Ventral surface light gray with sooty-
brown blotches scattered over disc, pelvic fins and tail. Tip of anterior lobes of
pelvic fins white. Large female paratype has a dark gray ventral surface. In life,
dorsal surface dark brown-black with pale yellow ocelli (Heemstra, pers. comm.).

VOLUME 95, NUMBER 3 447

sl
rh
sh
st
sp

Fig. 5. Right clasper of R. heemstrai RUSI 13812 (Holotype), partially expanded to show com-
ponents. fn—funnel, rh—rhipidion, sh—shield, sl—slit, sp—spike, st—sentinel.

Etymology.—Named after Phillip C. Heemstra (RUSI) who furnished us with
five of the specimens of the new species and who has been extremely cooperative
in furnishing us with elasmobranch material from South Africa.

Remarks.—Raja heemstrai most closely resembles Raja (Okamejei) powelli
Alcock, 1898, which has been reported from the Gulf of Martaban (Burma) and
the Gulf of Aden. Both species have rhomboid-shaped discs, rather long pointed
snouts,-and interspace between dorsal fins equal to or greater than the base of
the first dorsal fin, and thus differ from the other Okamejei species which have
less rhomboid, more rounded discs, shorter snouts, and an interdorsal space
which is less than the base of the first dorsal fin (except R. (O.) olseni Bigelow
and Schroeder, 1951, which has an interdorsal space equal to base of first dorsal
fin).

Raja heemstrai and R. powelli can be distinguished from each other by several

448 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Neurocranium of R. powelli BMNH 1939.5.24:17, 347 mm TL, male. a. Dorsal view, b.
Lateral view. Abbreviations as in Fig. 3.

proportional measurements, number of tooth rows in the upper jaw, coloration,
and structure of the neurocranium and scapulocoracoids. In R. poweilli the orbit
is less than the interorbital distance (greater in R. heemstrai), the preoral snout
length is about two times the mouth width (2% times in R. heemstrai) and the
distance between the first gill slits and the fifth gill slits is greater in R. powelli
than R. heemstrai (Table 1). Raja powelli possesses over 70 rows of teeth in the
upper jaw (31-35 in R. heemstrai) and a single ocellus at the base of each pectoral
fin (numerous ocelli in R. heemstrai). The neurocranium of R. powelli possesses
more expanded nasal capsules, broader interorbital distance, shorter rostral ap-
pendices, and a more triangular anterior fontanelle than R. heemstrai (Fig. 6,
Table 2). The scapulocoracoids of R. powelli are more anteroposteriorly expand-
ed but less expanded between the procondyle and mesocondyle than in R. heem-
strai and possess multiple postventral foramina rather than an expanded post-
ventral fenestra (Fig. 4b, Table 2).

Stehmann (1976) tentatively placed Raja philipi Lloyd, 1906, in Okamejei al-
though the holotype and only known specimen of this species from the Gulf of
Aden is lost. However, the validity of R. philipi is suspect. Norman (1939) syn-
onymized R. philipi and a species of Raja mentioned but not described as new
by Annandale (1909) with R. powelli. Stehmann (1976) considered R. powelli and
R. philipi distinct because of differences in the shape of the jaws, disc and nasal
curtains. Several of the distinctions noted by Stehmann, which are based on the

VOLUME 95, NUMBER 3 449

original illustrations of the two species plus the two specimens of R. powelli from
the Gulf of Aden (BMNH 1939.5.24:17—18), may be due to the state of preser-
vation of the holotype of R. philipi. From Lloyd’s illustrations (1908, Plates 40,
41) it appears that the jaws are partially protruded and that the pectoral fins are
fixed in an unnatural position, 1.e. the radials of the pectoral fins appear to be
bent caudally along their lateral extremities and creases are evident in the flesh
over the lateral aspect of the disc. Stehmann also justified recognition of both
species on the basis of capture localities of the specimens. He stated that the
type localities were the Gulf of Martaban, off Burma (R. powelli); Trevandrum,
off the southwest tip of India (Annandale’s specimen); and the Gulf of Aden (R.
philipi). However, the two BMNH specimens of R. powelli, examined by Steh-
mann and by us, were captured in the Gulf of Aden, the type locality of R. philipi.
It thus appears that R. powelli occurs along the entire northern Indian Ocean,
suggesting that R. powelli, R. philipi and R. sp. of Annandale are synonymous.
However, the status of R. philipi will remain uncertain until additional specimens
are available for study.

Raja (Okamejei) most closely resembles R. (Dipturus). According to Ishiyama
(1958, 1967) and Stehmann (1976) the two subgenera are distinguished by the
following characters: |. in Okamejei, snout about one-half head length (greater
than one-half head length in Dipturus); 2. disc width 1.2—1.5 times tail length
- (1.5-1.8 times tail length); 3. more than one nuchal thorn (generally one or no
nuchal thorns); 4. generally males with at least three rows of thorns on tail (males
with a single row of thorns); 5. small to medium sized species (large to very large
species); 6. proximal section of accessory terminal | cartilage of clasper distinctly
biramous (proximal section of accessory terminal | cartilage not distinctly bira-
mous); 7. anterior fontanelle of neurocranium with a distinct anterior margin
(anterior fontanelle without a distinct anterior margin). In our opinion only one
of these distinctions, the condition of the anterior fontanelle, is definitive. Snout
length and tail length ratios of Okamejei and Dipturus form a more or less con-
tinuous series (Bigelow and Schroeder 1953, 1958, 1962; Wallace 1967; Hulley
1970). Raja (Okamejei) olseni Bigelow and Schroeder, 1951 lacks a nuchal thorn
and R. (D.) laevis Mitchill, 1817 possesses three thorn rows on the tail. Okamejei
species are small to medium sized and Dipturus species are large to very large,
but the significance of this dichotomy is unknown. The proximal section of the
accessory terminal | cartilage in R. (D.) lanceorostrata is distinctly biramous. The
structure of the anterior fontanelle appears to be consistently distinct between
the two subgenera but worldwide comparisons of species of both subgenera are
needed to determine the validity and interrelationships of these two taxa.

Cruriaja andamanica (Lloyd, 1909)

The second known specimen of Cruriraja andamancia (ZMA 113.400) was
captured off Tanzania with one of the paratypes of R. heemstrai, ZMA 113.399.
This specimen closely resembles Lloyd’s (1909) illustration of the holotype (PI.
46, Fig. 2) and agrees with the proportional measurements listed by Stehmann
(1976) despite the fact that the holotype, at the time it was measured, was in poor
condition and in three pieces. The holotype differs from our specimen in pos-
sessing a shorter spiracle and greater distances between the first gill slits and

450 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

between the fifth gill slits (Table 1). Our specimen, like several other species of
Cruriraja, possesses oronasal pits similar to these described in Pseudoraja fis-
cheri (Bigelow and Schroeder, 1954), Breviraja, and Neoraja (McEachran and
Compagno, 1982). Cruriraja andamanica thus occurs in both the western and
eastern tropical Indian Ocean. Coloration of the two specimens is similar except
that the ventral surface of our specimen is grayish pink rather than slate gray.

Acknowledgments

We wish to thank Philip C. Heemstra (RUSI), Han Nijssen and Vertus van
Tuyl (ZMA), and Alwyne C. Wheeler and Peter J. P. Whitehead (BMNH) for
furnishing the specimens for this study. Thanks are also extended to Steven
Branstetter, Richard E. Matheson, Michael J. McCoid and Edward O. Murdy for
suggestions which improved the manuscript. This study was supported in part by
the National Science Foundation, Grant No. DEB 78-11217.

Literature Cited

Annandale, N. 1909. Report on the fishes taken by the Bengal fisheries steamer “‘Goldern Crown.”’
Part I. Batoidei.mMemoirs of the Indian Museum, 2: 1-60.

Bigelow, H. B., and W. C. Schroeder. 1953. Fishes of the western North Atlantic: sawfishes,

guitarfishes, skates and rays, and chimaeroids.—Memoir One, Sears Foundation of Marine

Research, 1(2):588 pp.

, and 1958. Four new rajoids from the Gulf of Mexico.—Bulletin of the Museum of

Comparative Zoology, 119:201-—233.

, and 1962. New and little known batoid fishes from the western Atlantic.—Bulletin

of the Museum of Comparative Zoology, 128:162—244.

Hulley, P. A. 1970. An investigation of the Rajidae of the west and south coasts of southern
Africa.—Annals of the South African Museum, 55:151—220.

Ishiyama, R. 1958. Studies on the rajid fishes (Rajidae) found in the waters around Japan.—Journal
of the Shimonoseki College of Fisheries, 7: 1-394.

———. 1967. Fauna Japonica. Rajidae (Pisces)—Tokyo Electrical Engineering College Press, To-
kyo. 84 pp.

Lloyd, R. E. 1908. Illustrations of the zoology of the Investigator. Fishes. pt. 9-10, pls. 39-50.

Calcutta.

1909. A description of the deep-sea fish caught by the R.I.M.S. ship ‘‘Investigator’’ since
the year 1900, with supposed evidence of mutation in Malthopsis.—Memoirs of the Indian
Museum, 2:139-180.

McEachran, J. D., and L. J. V. Compagno. 1979. A further description of Gurgesiella furvescens
with comments on the interrelationships of Gurgesiellidae and Pseudorajidae (Pisces, Rajoi-
dei).—Bulletin of Marine Science, 29:530-553.

, and 1982. Interrelationships of and within Breviraja based on anatomical structures
(Pisces, Rajoidei).—Bulletin of Marine Science, 32:399-425.

, and J. D. Fechhelm. 1982. A new species of skate from Western Australia with comments
on the status of Pavoraja Whitley, 1939 (Chondrichthyes, Rajiformes).—Proceedings of the
Biological Society of Washington, 95:1-12.

Norman, J. R. 1939. Fishes.—Scientific Report of the John Murray Expeditions, 1933-1934, 10):
1-116.

Stehmann, M. 1976. Revision des Rajoiden—Arten des nordlichen Indischen Ozean und Indopazifik
(Elasmobranchii, Batoidea, Rajiformes).—Beaufortia, 24(315):133-175.

Wallace, J. H. 1967. The batoid fishes of the east coast of southern Africa. Part III: Skates and
electric rays.—Oceanographic Research Institute, Investigation Reports, Durban, 17: 1-62.

(JDM) Department of Wildlife and Fisheries Sciences, and Department of
Oceanography, (JDF) Department of Wildlife and Fisheries Sciences, Texas A&M
University, College Station, Texas 77843.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 451-457

RELATIONSHIPS OF THE AFRICAN KILLIFISH
GENUS FOERSCHICHTHYS (TELEOSTEI:
CY PRINODONTIFORMES: APLOCHEILIDAE)

Lynne R. Parenti

Abstract.—The genus Foerschichthys was named by Scheel and Romand (1981)
for Aplocheilichthys flavipinnis Meinken, a diminutive species of killifish from
the Niger Delta. The relationship of this species to other killifishes has been
debated since its description. An osteological and external morphological ex-
amination reveals that Foerschichthys flavipinnis 1) is an aplocheiloid killifish; 2)
is a member of the solely Old World family Aplocheilidae; and 3) is more closely
related to the group including Aplocheilus, Pachypanchax and Epiplatys than it
is to the group including Aphyosemion, Fundulopanchax and Nothobranchius.
Foerschichthys is distinguished from all other aplocheiloid killifishes by the de-
rived placement of the dorsal fin entirely posterior to the anal fin.

Meinken (1929) reported the existence of a new, diminutive killifish species
(family Cyprinodontidae sensu Myers 1955) from the Niger Delta, referring to it
as ““Panchax species nov. der Zwergpanchax,’’ the dwarf or pygmy Panchax.
As used by Meinken (1929), Panchax was a general reference for Old World
killifishes of the subfamily Rivulinae Myers (=suborder Aplocheiloidei as defined
by Parenti 1981).

Meinken (1932) formally described the new dwarf Panchax as Aplocheilichthys
flavipinnis, thus changing his tentative suprageneric placement of the species from
the aplocheiloid killifishes (suborder Aplocheiloidei) to the relatively unrelated
aplocheilichthyine killifishes (suborder Cyprinodontoidei, family Poeciliidae,
subfamily Aplocheilichthyinae), more commonly known as the lampeyes or the
procatopines (see Parenti 1981). Primarily because the aplocheiloids and aplo-
cheilichthyines comprise the subsaharan African killifish fauna, they have often
been discussed as close relatives (e.g., Ahl 1924, 1928; Huber 1979; Scheel 1968;
Scheel and Romand 1981).

Scheel (1968) claimed that Aplocheilichthys flavipinnis possessed a greater
number of aplocheiloid than aplocheilichthyine characters, and suggested that it
be referred to as ‘“‘Aphyosemion’’ flavipinnis until a more precise statement of
relationship of the species could be made. (Aphyosemion is a large genus of
primarily West African aplocheiloids.) Scheel believed that the species was more
closely related to Aphyosemion than to Epiplatys, which, at that time, was the
other African aplocheiloid genus containing several diminutive forms.

Scheel and Romand (1981) compared A. flavipinnis to four species of diminutive
killifishes, the African Adamas formosus Huber, Epiplatys duboisi Poll, and E.
annulatus (Boulenger), and the South American Fluviphylax pygmaeus (Myers
and Carvalho), all of which they considered to be aplocheiloids. Because A.
flavipinnis differs from all four species, they named a new genus, Foerschichthys,
for the species. They were unable to make a statement concerning the relationship

452 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

of the genus other than its being aplocheiloid, and supported their taxonomic
decision by stating that (p. 30): “*. . . we find ourselves unable to place A. fla-
vipinnis in any known rivulin genus.”’

Parenti (1981) reclassified the killifishes, the order Cyprinodontiformes, into
two suborders, the Aplocheiloidei and the Cyprinodontoidei, defining each order
as monophyletic (sensu Hennig 1966) using derived, primarily osteological, char-
acters. Fluviphylax pygmaeus was classified as a cyprinodontoid (in the family
Poeciliidae, subfamily Fluviphylacinae), whereas the other species compared by
Scheel and Romand (1981) were classified as aplocheiloids. The relationship of
Fluviphylax to the aplocheilichthyines and the viviparous poeciliids was pre-
sented as an unresolved trichotomy by Parenti (1981), and that genus should not
be considered an aplocheiloid as it was by Scheel and Romand (1981).

The present investigation was prompted by my own observations and those of
Scheel (1968) and Scheel and Romand (1981) that Foerschichthys shares with
Fluviphylax pygmaeus the derived character of the placement of the dorsal fin
posterior to the anal fin.

The objectives of the present study were to determine 1) if there are uniquely
derived characters to define Foerschichthys ; 2) if the genus is more closely related
to the aplocheiloids or to the cyprinodontoids (including the aplocheilichthyines);
and 3) to what genus or group of genera it is most closely related. The possibility
that Foerschichthys is a close relative of Fluviphylax was considered initially and
then rejected after an examination of material revealed that Foerschichthys is an
aplocheiloid killifish.

Discussion

Aplocheiloid and aplocheilichthyine killifishes superficially resemble each other
and have often been confused in the literature. The suborder Aplocheiloidei (order
Cyprinodontiformes) is defined as monophyletic by nine derived characters (Pa-
renti 1981:374): 1) attached orbital rim; 2) cartilaginous mesethmoid; 3) close-set
pelvic girdles; 4) a broad anterior end of basihyal; 5) a narrow and twisted preor-
bital bone; 6) tubular anterior naris; 7) reduced cephalic sensory pore pattern; 8)
males more elaborately pigmented than females; and 9) posterior extension of
vomer dorsal to anterior ramus of parasphenoid. The genera Foerschichthys,
Adamas, Aphyosemion and Epiplatys are members of the suborder Aplochei-
loidei.

Although some of these characters appear in the suborder Cyprinodontoidei,
their occurrence in that suborder has been most parsimoniously assessed as ho-
moplasious by Parenti (1981). Twelve derived characters were given to define the
suborder Cyprinodontoidei as monophyletic (Parenti 1981:403-404). These in-
clude two rather than three ossified basibranchials, and the absence of the dorsal
hypohyal, the metapterygoid, the first dorsal fin ray, as well as several upper jaw
ligaments. Aplocheilichthyine killifishes, as well as Fluviphylax pygmaeus, are
members of the suborder Cyprinodontoidel.

Within the Aplocheiloidei, there are two families, the Old World Aplocheilidae
and the New World Rivulidae. The Aplocheilidae is defined as monophyletic by
four derived characters (Parenti 1981:387): 1) supracleithrum fused to posttem-
poral; 2) basihyal a small triangular bone capped by a large wedge of cartilage;

VOLUME 95, NUMBER 3 453
Aplocheilus
Pachypanchax

Epiplatys

Foerschichthys

Adamas

Aphyosemion

Fundulopanchax

Nothobranchius

Fig. |. Cladogram of relationships of the genera of the family Aplocheilidae. Derived characters
at each lettered node are as below. For characters defining other nodes, and a discussion of relation-
ships of genera and their included subgenera, see Parenti (1981). a. Posttemporal fused to supra-
cleithrum; reduction of ossified section of basihyal to a small, triangular wedge; interarcual cartilage
attaches directly to the articulation point of the second pharyngobranchial; premaxillary processes
tapered posteriorly. b. Reduction of the uncinate process on the fourth epibranchial; premaxillary
ascending processes expanded. c. Expanded coronoid process on the dentary; bifurcate dorsal hypural
plate in juveniles and some adults; absence of the uncinate process of the fourth epibranchial. d. Bifid
epipleural ribs; attenuate posterior extension of the vomer.

3) premaxillary ascending processes tapered posteriorly; and 4) interarcual car-
tilage attaching directly to articulation point of second pharyngobranchial. Foer-
schichthys, Adamas, Aphyosemion and Epiplatys are members of the Aplochei-
lidae.

The family was divided informally into two groups that were not named as
subfamilies by Parenti (1981) because the distinction among species of the most
primitive genera of each group is problematic. However, Foerschichthys pos-
sesses two of the defining characters of the Aplocheilus-Pachypanchax-Epiplatys
group (Parenti 1981:395). The premaxillary ascending processes are relatively
expanded, although not as greatly as in Aplocheilus panchax (Parenti 1981:359,
Fig. 4a), rather than being attenuate as in Aphyosemion petersi (Fig. 4c). Also,
in Aplocheilus panchax (Parenti 1981:388, Fig. 24a) there is no uncinate process
on the fourth epibranchial, and therefore the fourth epibranchial is not in contact
with the third. In Foerschichthys the uncinate process of the fourth epibranchial
is reduced, with a cartilaginous cap for articulation with the uncinate process of
the third epibranchial. The relationships of the major groups of aplocheiloid gen-
era are summarized in Fig. 1. A clearer statement of the relationships among all
nominal aplocheiloid genera and subgenera awaits a revision of all included
species.

454 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Scheel and Romand (1981) defined Foerschichthys phenetically, stating that it
differed from other African aplocheiloids (of which group they considered Flu-
viphylax to be a member) by a combination of three characters: 1) pigmentation
pattern; 2) karyotype; and 3) origin of the dorsal fin. Only the state of the last
character was analyzed in terms of being primitive or derived, and it is the only
derived character by which the genus is defined.

The pigmentation pattern is characterized by a narrow, black median line, and
a similar black line along the base of the anal fin. Such a pigmentation pattern
characterizes many atherinomorph fishes (the larger group to which cyprinodon-
tiforms belong) and is commonly found in aplocheilichthyines among the cyprin-
odontiforms. Because of its wide distribution, the pigmentation pattern should
initially be assessed as primitive for atherinomorph fishes. However, considering
that, as listed above, a defining character of the suborder Aplocheiloidei is that
males are more elaborately pigmented than females, the much less elaborate
pattern of Foerschichthys is perhaps secondarily derived. As stated by Scheel
and Romand (1981:23): ‘‘Such narrow black lines occur in almost all juveniles
and adult procatopodin [aplocheilichthyine] species but are very rare in rivulin
[aplocheiloid] fish; they have been observed only in Epiplatys annulatus and in
the two Asian fishes Aplocheilus blocki (Arnold, 1911) and A. panchax (Hamilton-
Buchanan, 1822), but in these fishes the lines disappear with age.’’ Thus, if the
pattern is considered secondarily derived, it further supports the alignment of
Foerschichthys with the Aplocheilus-Pachypanchax-Epiplatys group.

The general number of haploid chromosomes in teleost fish is 24. Fishes of the
Aphyosemion-Nothobranchius group typically have a very low haploid chromo-
some number; the lowest number reported is nine (Scheel 1968). Foerschichthys
flavipinnis has a haploid chromosome number of 20 which Scheel and Romand
(1981) point out as a character that distinguishes it from the other diminutive
aplocheiloids. However, it is not a unique karyotype for aplocheiloid fishes. Fur-
thermore, given that the haploid chromosome number ranges from nine to 25 in
cyprinodontiforms, the karyotype of Foerschichthys may simply cepresem a Stage
in a transition series from a high to a low chromosome number.

The only derived character defining Foerschichthys within the aplocheiloids is
the placement of the dorsal fin posterior to the anal fin. Fluviphylax pygmaeus
shares this character but is closely related to the aplocheilichthyine cyprinodon-
toids. The occurrence of this character in both genera indicates that it is inde-
pendently and secondarily derived.

The description that follows is based on personal observation as well as data
from Meinken (1932), Scheel (1968) and Scheel and Romand (1981). The char-
acters and their order are similar to those given for other cyprinodontiform genera
by Parenti (1981).

Foerschichthys Scheel and Romand

Foerschichthys Scheel and Romand, 1981:30, type by original designation
Aplocheilichthys flavipinnis Meinken.

Diagnosis.—Distinguished from all other aplocheiloid killifishes by having the
dorsal fin situated entirely posterior to the anal fin.

VOLUME 95, NUMBER 3 455

Foerschichthys flavipinnis (Meinken)

‘‘Panchax species nov.’’ Meinken, 1929:385, aquarium description.

Aplocheilichthys flavipinnis Meinken, 1932:54, original description, Nigeria, La-
gos.—Kaden, 1967:280, aquarium description, color pattern.—Foersch, 1968:
142-143, aquarium description.—Scheel, 1968:211—214, discussion of relation-
ships.

‘‘Aphyosemion’”’ flavipinnis Scheel, 1968:214, tentative referral of this species to
Aphyosemion.

Foerschichthys flavipinnis Scheel and Romand, 1981:30, proposal of a new genus
with A. flavipinnis as its type. Combination **Foerschichthys flavipinnis’’ used
only in figures on pages 24 and 25, and table on page 26.

Diagnosis.—Same as for genus.

Description.—Anal: iv, 10; Dorsal: 11, 6; Pelvic: 6; Pectoral: 13; Vertebrae: 13
+ 17; Scales in lateral series: 30-31.

First pleural rib on parapophysis of second vertebra; parapophysis not reduced;
no pleural rib on the first hemal spine; hypural plates fused into an were fan.
Epipleural ribs not bifid.

Anal fin not modified into a gonopodium; anal fin musculature unmodified; first
proximal radial present; middle anal radials present.
~ One dorsal fin ray articulating with each of first two dorsal radials; dorsal fin
origin posterior to anal fin.

Autopterotic fossa normal; lateral ethmoid not expanded medially, not reaching
parasphenoid; parasphenoid not expanded anteriorly; weakly formed supraoccip-
ital and epiotic processes; neural spine on first vertebra; first vertebra articulates
with skull via basioccipital and exoccipital condyles; supraoccipital excluded from
formation of foramen magnum; parietals present; nasals not expanded medially.

Mesethemoid cartilaginous; medial process of pelvic fin base and ischial pro-
cess reduced; interarcual cartilage attached directly to articulation point of second
pharyngobranchial; basihyal broad anteriorly, small triangular ossification pos-
teriorly; no tooth patches on second and third hypobranchials; few teeth on fourth
ceratobranchials; dorsal and ventral hypohyal present; anterior extension of an-
terior ceratohyal ventral to hypohyals; reduced uncinate process on fourth epi-
branchial articulates with that of third; first epibranchial narrow at its base. In-
terhyal ossified; three ossified basibranchials. Vomer with posterior extension
dorsal to parasphenoid.

Preorbital bone narrow and twisted, carrying distinct sensory canal; dermo-
sphenotic and preopercle with distinct sensory canal; pectoral girdle lowset; first
postcleithrum present; posttemporal straight, with ligamentous ventral limb; post-
temporal fused to supracleithrum.

Vomer present, edentulous; medial ramus of maxilla twisted with no pro-
nounced dorsal process, ventral process gently curved toward and abutting rostral
cartilage, lateral ramus narrow.

Premaxillary ascending processes flat and broad, slightly tapered posteriorly,
not overlapping in the midline; rostral cartilage large; lateral ramus of premaxilla
with alveolar process not indented posteriorly. Ligament extending from ventral
processes of maxillae to middle of rostral cartilage; ethmomaxillary ligament
present; meniscus present between premaxilla and maxilla.

456 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Dentary not expanded medially, not robust; coronoid process on dentary not
overlapping that of angulo-articular; retroarticular not elongate. Autopalatine with
straight head, ventral process not elongate, not reaching quadrate; metapterygoid
reduced.

Orbital rim attached; anterior naris tubular; supraorbital sensory pores reduced
to a series of neuromasts situated in open troughs; preopercular and mandibular
canals open troughs; two preorbital pores.

Males larger than females. Pigmentation pattern: thin median black line, thin
black line on base of anal, males with scales above median black line edged in
red; dorsal and anal fins yellow to orange, males with light blue fin edges; no spot
at anterior base of dorsal fin; throat bars present.

No fatty predorsal ridge; caudal fin scaled for approximately one-third its length;
caudal fin with a single, median posterior point; swimbladder extends posteriorly
to first hemal spine.

Distribution.—Known from following Nigerian localities: Ibefun, south of Te-
bu-Ode; Ughelli, east of Warri of the Niger Delta; Aduna, about 67 km south of
Isela-Ubu; Auna, between Ijebu-Ode and Shagamu; Aiyetoto (Scheel 1968).

Remarks.—Meinken (1932:53) stated that the types of Aplocheilichthys flavi-
pinnis were to be deposited in the Berliner Zoologische Museum, but a search of
the collection by Dr. H.-J. Paepke failed to locate them. Nevertheless, a neotype
will not be designated at this time, following the recommendations of the Inter-
national Code of Zoological Nomenclature, because it is possible that the type
material is mislabelled in the Berlin Museum or is at another institution.

Material Examined

Foerschichthys flavipinnis: Nigeria: Niger Delta at Ughelli: MCZ 49881 (4 spec.,
1 specimen counterstained with alcian blue and alizarin red S).

Fluviphylax pygmaeus: Brazil: Manaus: MCZ 46712 (21 spec.); MCZ 46713 (64
spec., 3 spec. counterstained); MCZ 46714 (11 spec.); MCZ 49958 (14 spec., 4
spec. cleared and alizarin stained); Lago Hyanuary: MCZ 41367 (5 spec.); Rio
Madeira at Borba: SU 50196 (3 spec.) Paratypes. :

Adamas formosus: Zaire: Village of Ntokon near the banks of the Likouala-
Mossaka: MNHN 1979-199 (1 spec.) Holotype. 3

Epiplatys annulatus: Sierra Leone: BMNH 1914.12.9:5—6 (2 spec.). Syntypes.

Additional comparative data were taken from Parenti (1981).

Summary

The monotypic African killifish genus Foerschichthys was named by Scheel
and Romand (1981) for the diminutive Aplocheilichthys flavipinnis Meinken be-
cause they could not place it in any of the known tropical killifish genera with
dwarf forms (Aphyosemion, Adamas, and Epiplatys of Africa and Fluviphylax of
South America). Foerschichthys is distinguished from all other aplocheiloid gen-
era by having the dorsal fin situated entirely posterior to the anal fin.

An examination of osteological and other anatomical features of Foerschichthys
flavipinnis reveals that it does not possess the derived features of the suborder
Cyprinodontoidei, the group in which Fluviphylax is placed (Parenti 1981), but
that it possesses the derived features of the suborder Aplocheiloidei. Foerschich-

VOLUME 95, NUMBER 3 457

thys may be considered a primitive member of the Aplocheilus-Pachypanchax-
Epiplatys group of the family Aplocheilidae.

Acknowledgments

I thank H.-J. Paepke, Museum fur Naturkunde der Humboldt-Universitat, Ber-
lin, for searching through the collection for the type specimens of Aplocheilich-
thys flavipinnis Meinken.

The following persons lent or allowed me to examine specimens in their care:
W. L. Fink (Museum of Comparative Zoology, Cambridge, MCZ); W. N. Esch-
meyer (Stanford University, SU, now at the California Academy of Sciences,
San Francisco); M. L. Bauchot (Muséum National d’Histoire Naturelle, Paris,
MNHN); and, P. H. Greenwood (British Museum [Natural History], London,
BMNH).

This research was initiated during the tenure of a Smithsonian Postdoctoral
Fellowship, National Museum of Natural History, and completed during a NATO
Postdoctoral Fellowship at the British Museum (Natural History). The manuscript
benefitted greatly from the comments of a reviewer, and discussions with Jean
H. Huber about African killifish. |

Literature Cited

Ahl, E. 1924. Zur Systematik der altweltlichen Zahnkarpfen der Unterfamilie Fundulinae.—Zool-
ogischer Anzeiger 60:49-55.
1928. Beitrage zur Systematik der afrikanischen Zahnkarpfen.—Zoologischer Anzeiger 79:

113-123.

Foersch, W. 1968. Aplocheilichthys flavipinnis Meinken, 1932, der gelbflossige Leuchtaugenfisch.—
Die Aquarien- und Terrarien-Zeitschnft. (DATZ). Jahrgang 21(4): 142-143.

Hennig, W. 1966. Phylogenetic systematics.—University of Illinois Press, Urbana. 263 pp.

Huber, J. H. 1979. Cyprinodontides de la cuvette congolaise (Adamas formosus n. gen., n. sp. et
nouvelle description de Aphyosemion splendidum).—Revue Frangaise d’ Aquariologie Herpé-
tologie 6(1):5—10.

Kaden, D. 1967. Ein ganz Neuer aus Afrika.—Aquarien-Terrarien [German Democratic Republic]
14:280.

Meinken, H. L. 1929. Vorlaufige Mitteilungen tiber zwei neue, noch unbestimmte Zahnkarpfen aus

Westafrika.—Blatter fir Aquarien und Terrarienkunde, 40:385-389.

1932. Uber einige neue Zahnkarpfen aus dem tropischen Westafrika.—Blatter fiir Aquarien

und Terrarienkunde, 43:53-58.

Myers, G. S. 1955. Notes on the classification and names of cyprinodont fishes.—Tropical Fish
Magazine 4(4):7.

Parenti, L. R. 1981. A phylogenetic and biogeographic analysis of cyprinodontiform fishes (Tel-
eostei, Atherinomorpha).—Bulletin of the American Museum of Natural History 168(4):335—
357?

Scheel, J. J. 1968. Rivulins of the Old World.—T. F. H. Publications, Inc., New Jersey. 473 pp.

, and R. Romand. 1981. A new genus of rivulin fish from tropical Africa (Pisces, Cyprino-

dontidae).—Tropical Fish Hobbyist 29(11):22-30.

NATO Postdoctoral Fellow, Department of Zoology, British Museum (Natural
History), London SW7 5BD, England.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 458-477

A NEW XYSTODESMID MILLIPED GENUS AND THREE
NEW SPECIES FROM THE EASTERN BLUE RIDGE
MOUNTAINS OF NORTH CAROLINA |
(POLYDESMIDA)

Rowland M. Shelley

Abstract.—The new xystodesmid milliped genus Prionogonus is proposed for
three new species—iaerens, divaricatus, and thrinax—in the eastern Blue Ridge
Mountains and Blue Ridge escarpment of southwestern North Carolina. A row
of blunt or sharply pointed spurs along the lateral side of the basal zone of the
gonopodal acropodite, not found in any other apheloriine genus, characterizes
Prionogonus, which is also distinguished by the absence of a distal zone and
apical curve from the acropodite and by the presence of a solenomerite. Specific
differences include the length of the row of spurs, the overall configuration and
apical features of the acropodite, and the position and configuration of the solen-
omerite. The species are allopatric but occur close together in limited ranges.
The distribution of haerens, much larger than that of either of the others, covers
over half the generic range, which is the smallest in the tribe Apheloriini. Prio-
nogonus is a “‘sigmoid’’ genus and may share common ancestry with Sigmoria
stibarophalla Shelley. The species haerens and divaricatus evolved from a com-
mon ancestor, but thrinax is more distant.

This contribution concerns a group of ‘‘sigmoid’’ xystodesmid millipeds that
inhabits a small area in the southern Blue Ridge Mountains and whose existence
has been known for several years. Diagnoses were deferred until taxonomic con-
fusion among the established *‘sigmoid’’ genera—Sigmoria, Sigiria, and Fallo-
ria—could be resolved, because I did not want to risk having to synonymize a
new genus on the basis of revised concepts of existing genera. I have now dem-
onstrated that Sigiria and Falloria are synonyms of Sigmoria, and broadened the
concept of this taxon to encompass a diversity of forms whose gonopods conform
to a particular pattern (Shelley 1981la). The gonopods of the xystodesmids dis-
cussed herein differ significantly from this pattern, leaving no doubt that they
represent a new apheloriine genus.

I defined Sigmoria to include forms in which the gonopodal acropodite curves
through one or more vertical planes and consists of three major sections labeled
‘‘basal zone,”’ ‘‘peak,’’ and ‘‘distal zone’’ (Shelley 1981a) (see gonopod termi-
nology section under taxonomic characters). The basal zone and peak are sepa-
rated by a broad or sharp “‘anterior bend,’’ and the acropodite usually displays
an “‘apical curve,’’ which is formed by the distal zone. Every species of Sigmoria
except one possesses a flange on the medial face of the acropodite, either on the
proximal portion of the peak or on the proximal portion of the distal zone. Al-
though the tip of the acropodite is variable, there is no solenomerite; the acro-
podite tapers to a single apical termination, on which the prostatic groove opens.
Two species differ from aspects of this pattern. Sigmoria nantahalae Hoffman

VOLUME 95, NUMBER 3 459

lacks a medial flange, and the distal zone and apical curve are absent from S.
truncata Shelley. Both of these species otherwise conform to the generic defi-
nition and are easily accommodated by Sigmoria. Because the three species
described herein have gonopods that differ markedly from the above plan, broad-
ening the concept of Sigmoria to include them would destroy the homogeneity
the genus now possesses. These new species all lack the distal zone and apical
curve, and the acropodites terminate at the distal extremities of the peaks. Thus,
the consistent absence of the distal part of the acropodite cannot be interpreted
as an individual secondary modification of the basic Sigmoria plan, as is possible
in the case of S. truncata, but instead represents an entirely different gonopodal
pattern. Furthermore, these three species all possess a distinct solenomerite. In
one species the acropodite ends in three acute projections, with the central one
carrying the prostatic groove, and in the others the solenomerite is located be-
neath the distal extremity of the peak so as to form a separate projection from
the ending of the acropodite proper. All three species exhibit one Sigmoria trait—
a medial flange on the proximal portion of the peak. All three additionally display
a distal tooth on the flange, which may be homologous to the tooth of some
species of Sigmoria, particularly those of the latior species group. However, two
of the new species also possess a lateral flange on be peak opposite the medial
flange, a feature not shared with species of Sigmoria. Finally, all three new
species possess a row of distinct spurs on the lateral side of the basal zone, a
feature unique to “‘sigmoid’’ xystodesmids. The tubercles of S. tuberosa Shelley
are neither homologous nor analogous to the spurs, since they are much smaller
and are clustered basally.

Thus, these three new species represent a new “‘sigmoid’’ genus—Prionogo-
nus—whose name refers to the spurs on the basal zone. Its species are distin-
guished by the overall configuration and apical characteristics of the acropodite,
by the position and configuration of the solenomerite, and by the length of the
row of spurs. Either I or an assistant collected most of the material on which this
paper is based, and our specimens are deposited in the invertebrate collection of
the North Carolina State Museum of Natural History (NCSM). In the ensuing
species accounts the collector’s name is omitted for samples taken by me alone,
and the NCSM invertebrate catalog number is indicated in parentheses after
appropriate locality citations. Four samples of the most widely distributed species
were discovered in the collection of the Museum of Comparative Zoology (MCZ)
and the private holdings of Richard L. Hoffman (RLH). Paratypes of all three
species were deposited in the Florida State Collection of Arthropods (FSCA),
Gainesville.

6

Taxonomic Characters

The taxonomic characters of Prionogonus are found exclusively on the gono-
pods. Color pattern is of no taxonomic value because sympatric species of Sig-
moria also exhibit the red paranota and metatergal stripes seen in Prionogonus.
The length and configuration of the process of the 4th sternum also differ in each
species (longer than the adjacent coxal widths in haerens, slightly shorter than
the coxae and deeply divided apically in divaricatus, and much shorter than the
coxae and with four long apical setae in thrinax), but it would be very risky to

460 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

base identifications on this structure alone. Differences between this process in
species of Prionogonus and sympatric species of Sigmoria are of insufficient
magnitude to allow for authentic determinations.

The most important and obvious diagnostic character of the gonopods of Prio-
nogonus is the row of spurs on the lateral side of the basal zone. No other
xystodesmid genus in the eastern United States possesses such a feature.

At the species level, the taxonomically useful characters are located solely on
the acropodites, as differences in the size and configuration of the prefemoral
processes are too insigifinicant to be reliable. The greatest utility of the spurs is
in distinguishing divaricatus from its congeners. In this species the spurs extend
around the anterior bend onto the peak and terminate at the base of the lateral
flange. The spurs on both haerens and thrinax are restricted to the basal zone;
they are smaller and fewer in number in thrinax, but this difference is of little
taxonomic use.

The overall configuration of the acropodite is an important taxonomic char-
acter. In thrinax the anterior bend is broad and poorly defined, and the acropodite
forms a smoothly continuous arc of broad diameter. In haerens and divaricatus
the anterior bend is sharp, approximately a right angle, and the configuration of
the acropodite resembles an inverted L. The peak of the acropodite is slightly
longer in divaricatus and hence overhangs the prefemoral process to a greater
degree than in haerens. Consequently, the relative proportions of the basal zone
and peak to the total length of the acropodite also differ in these two species.
Each region represents about half of the total acropodite length in divaricatus,
whereas the basal zone comprises about two-thirds, and the peak one-third, of
the length in haerens.

Apically on the acropodite, the three-pronged termination, the subspiniform
solenomerite directed away from the gonopod along the axis of the acropodite,
and the presence of the lateral process are all diagnostic for thrinax. In separating
haerens from divaricatus, fusion or segregation, respectively, of the acropodite
and the undersurface of the peak are the most important diagnostic features. Of
nearly equal taxonomic value, however, is the size of the tooth on the medial
flange. In divaricatus it is large and extends to the level of the distal extremity
of the peak, but in haerens it is reduced, vestigial, or absent, and terminates
proximal to the end of the peak when present.

Prionogonus, new genus

Type species.—Prionogonus haerens, new species.

Description.—A genus of moderate-size xystodesmids with the following char-
acteristics: |

Body composed of head and 20 segments in both sexes; size varying from
around 40—45 mm in length and 9.5—11.5 mm in width; W/L ratio similarly varying
from about 22.5—25.5%. Body essentially parallel-sided in midbody region, ta-
pering at both ends.

Color in life constant; all forms with red paranota and red connecting stripes
along caudal edges of metaterga, with additional red connecting stripe along an-
terior edge of collum.

VOLUME 95, NUMBER 3 46|

Head of normal appearance, smooth, polished. Epicranial suture thin but dis-
tinct, terminating in interantennal region, not apically bifid; interantennal isthmus
varying from around 1.5—2.0 mm; genae not margined laterally, with shallow
central impressions, ends broadly rounded and projecting slightly beyond adja-
cent cranial margins. Antennae moderately slender, relatively constant in length,
becoming progressively more hirsute distally, with 4 conical sensory cones on
ultimate article; no other sensory structures apparent. Facial setae reduced; epi-
cranial and interantennal absent, frontal and genal present or absent, clypeal and
labral present.

Terga smooth and polished, becoming moderately coriaceous in paranotal re-
gions, especially on anterior half of paranota. Collum broad but variable, ends
subequal to or extending slightly beyond those of following tergite. Paranota
moderately to strongly depressed but generally continuing slope of dorsum, an-
teriolateral corners rounded on all segments, caudolateral corners rounded on
anteriormost segments, becoming blunt in midbody region and progressively more
acute posteriorly. Peritremata flat and indistinct, especially in anterior third of
body, not strongly elevated above metazonal surface; ozopores located just cau-
dal to midlength, opening dorsad. Prozonites smaller than metazonites; strictures
moderately distinct, slightly costulate.

Caudal segments normal for family.

Sides of metazonites generally smooth, without grooves or impressions. Pre-
gonopodal sterna of males variously modified; that of segment 4 with medial
process of variable length, shorter to longer than widths of adjacent coxae, with
or without apical setae; sternum of segment 5 with two distinct or medially co-
alesced processes between 4th legs, usually slightly shorter than widths of adja-
cent coxae, with variable elevated areas or knobs between 5th legs; sternum of
segment 6 with minute processes between 6th legs and with shallow convex
recession between 7th legs to accommodate apical regions of acropodites when
body segments compressed; 7th legs occasionally set slightly farther apart than
6th. Postgonopodal sterna with bicruciform impressions on first few segments,
becoming flattened and plate-like posteriorly with small central impressions, with-
out noticeable lobes along caudal edges between posterior segmental legs. Gono-
pores on second pair of legs of males short, with round, apical knobs. Coxae with
varying distomedial projections beginning on midbody (postgonopodal) legs, usu-
ally low, blunt tubercles, occasionally more acute; prefemoral spines relatively
long and sharply pointed; tarsal claws hooked or bisinuate. Hypoproct broadly
rounded; paraprocts with margins strongly thickened.

Gonopodal aperture elliptical, indented and slightly narrower anteriolaterally,
sides flush with or elevated above metazonal surface. Gonopods in situ with
acropodites projecting ventrad from aperture, bending anteromediad and crossing
midline of aperture, curving smoothly over opposite side of aperture and termi-
nating either over aperture or slightly beyond anterior margin, either overlapping
each other in midline of aperture or curving in front of and behind each other.
Coxae moderate, without apophysis, connected by membrane only, no sternal
remnant. Prefemora moderate in size, with small but variable, apically blunt
prefemoral processes arising on anteriomedial sides. Acropodites moderately thick
and heavy, well sclerotized, curving through mostly a single vertical plane, bent

462 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

sharply anteriad (anterior bend) at % to % of total length or curving through
continuous arc of broad diameter, terminating distal to anterior bend in peak
region; portion between prefemur and anterior bend (basal zone) with row of
spurs along lateral margin; spurs variable in length, acuteness, and degree of
separation, terminating on basal zone or continuing around anterior bend onto
peak; anterior bend either sharp, 90°, and well defined, or broad and poorly
defined; portion distal to anterior bend (peak) varying from % to % of total
acropodite length, extending to or slightly beyond level of prefemoral process,
with flange of variable width and length on medial side, with or without similar
flange on lateral side; medial flange with slightly convex margin, usually with
small to moderate, subacute tooth at distal corner, tooth either extending to level
of distal extremity of peak or terminating short of this point, either directed away
from gonopod along axis of acropodite or downward toward coxa and subper-
pendicular to axis; lateral flange subequal to or slightly longer than medial flange,
usually subequal in width to medial flange, margin broadly wavy to nearly straight;
lateral terminal process present only when lateral flange absent, moderately long,
subacute apically; solenomerite either a bisinuate process fused to or narrowly
separated from distal extremity of peak, located beneath and directed laterad,
perpendicular to peak, or a long, subspiniform, apically recurved projection di-
rected away from gonopod along axis of acropodite, widely separated from lateral
process and tooth of medial flange forming trifurcate termination to acropodite.
Prostatic groove arising in pit on medial side of prefemur, crossing to lateral side
of acropodite on basal zone just distal to juncture with prefemur, running along
stem of acropodite on lateral side and opening terminally on tip of solenomerite.

Cyphopodal aperture short and very broad, encircling 2nd legs, sides elevated
above metazonal surface. Cyphopods in situ located lateral to 2nd legs, with
corner of receptacle and valves visible in aperture, valves directed caudad, ori-
ented dorsoventrally in body. Receptacle variable, moderate to large, located on
anterior and lateral sides of valves, cupped over ventral corner of valves in two
species, with or without noticeable lobes on anterior and lateral sides, surface
rugulose with distinct folds and ridges. Valves moderate in size, either subequal
or medial (inner) one slightly larger, surface generally smooth, finely granulate.
Operculum minute, located under free (dorsal) end of valves.

Distribution.—Eastern Blue Ridge Mountains and the Blue Ridge escarpment
of southwestern North Carolina, ranging from the southern edge of the Black
Mountains in McDowell and Buncombe counties to near the Green River on the
Henderson-Polk counties line. In the north-south direction the area extends some
26 miles from slightly north of I-40 near the towns of Black Mountain and Old
Fort to about four miles north of I-26 and the towns of Flat Rock and Saluda. In
the north, the east-west distribution extends from near Swannanoa to Old Fort,
a distance of about 13 miles, whereas in the south, the range narrows to a point
about four miles north of Saluda.

Species.—Three. One wide-ranging species occupies the northern half of the
range; the southern half is inhabited by all three species, the other two having
restricted ranges. One or two undiscovered species may exist in inaccessible parts
of the southern half of the range, as explained in the distribution section of the

paper.

VOLUME 95, NUMBER 3 463

Key to Species of Prionogonus (based on adult males)

—

. Acropodite with trifurcate termination, solenomerite directed away from
gonopod along axis of acropodite; acropodite a broad continuous curve,
anterior bend poorly defined; Polk and Henderson counties ...........

POM cate eee er ee ale acs oslo gies he bk oa es thrinax, new species
Acropodite termination blunt to subacute, with at most two terminal pro-
jections; solenomerite located beneath distal extremity of peak, directed
perpendicularly and laterad to peak; acropodite bent sharply at 4 to 1%
lenetheamiemormend well defined ......... cs. ...5205 555050005000.

. Solenomerite fused to undersurface of peak; spurs moderate in size, lo-
cated entirely on basal zone; McDowell, Buncombe, and Henderson
COMMUNES 0» ods S'S eee ne Ie a ea eet al haerens, new species
Solenomerite a separate bisinuate process, narrowly segregated from peak;
spurs moderate to large in size, extending around anterior bend to base
of lateral flange on peak; Rutherford, Henderson, and Polk counties

sc Es Se oc te ee see 2 1 oa HALVOTICGIUS,. NEW SPECIES

N
N

Prionogonus haerens, new species
Figs. 1-4

Type-specimens.—Male holotype (NCSM A988) and six male and five female
paratypes collected by R. M. Shelley, 12 July 1976, from Buncombe County, NC,
12.6 mi. SE Asheville, along US highway 74, 0.4 mi. W Henderson County line.
One male and two female paratypes taken by same collector on same date, 12.4
mi. SE Asheville, along US highway 74, 1.0 mi. W Henderson County line. Male
and female paratypes deposited in FSCA and RLH.

Diagnosis.—Characterized by fusion of solenomerite to undersurface of distal
extremity of peak; tooth of medial flange absent, vestigial, or moderate in size,
not projecting to level of distal extremity of peak; spurs moderate in size, oc-
curring only on basal zone.

Holotype.—Length 41.9 mm, maximum width 9.6 mm, W/L ratio 22.9%, depth/
width ratio 58.3%. Segmental widths as follows:

collum 6.4 mm 12th-14th 9.4
2nd 8.3 15th 9.0

3rd 9.0 16th 8.6

Ath 9.4 17th a>
Sth—11th 9.6 18th ey,

Color in life: paranota red; metaterga black with wide, red, transverse stripes
along caudal edges connecting paranotal spots; collum with red stripes along both
anterior and caudal margins.

Head capsule smooth, polished, width across genal apices 5.5 mm; interanten-
nal isthmus 1.4 mm, smooth; epicranial suture thin but distinct, terminating in
slight impression in interantennal region, not bifid. Antennae moderately long and
slender, reaching back to middle of paranota of third segment, becoming pro-
gressively more hirsute distally, with four terminal sensory cones, no other sen-

464 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-4. Prionogonus haerens: 1, Process of 4th sternum of holotype, caudal view (leg setation
omitted in all sternal drawings); 2, Gonopods in situ, ventral view of holotype; 3, Left gonopod of
holotype, medial view (setation omitted in all dissected gonopod drawings); 4, The same, lateral view.
Scale line for Fig. 2 = 1.00 mm; line for other Figs. = 0.75 mm for 1, 1.00 mm for 34.

sory structures evident, relative lengths of antennomeres 2 > 3 > 4 =
5 =6> 1>7, 1 subglobose, 2-6 clavate, 7 short and truncate. Genae not mar-
gined laterally, with distinct medial impressions, ends broadly rounded and pro-
jecting slightly beyond adjacent cranial margins. Facial setae as follows: epicran-
lal, interantennal, frontal, and genal absent, clypeal about 10-10, labral about 20-—
20.

Terga smooth, polished, becoming moderately coriaceous on paranota. Collum
broad, ends extending slightly beyond those of following tergite, caudal edge
relatively straight. Paranota moderately depressed, angled ventrad and continuing

VOLUME 95, NUMBER 3 465

slope of dorsum; anterior corners rounded; caudolateral corners rounded only on
first three tergites, blunt on segments 4-15, becoming acute and pointed on re-
maining segments. Peritremata shallow, indistinct on first eight segments; only
slightly elevated above metazonal surface thereafter. Ozopores located caudal to
midlength of peritremata, opening dorsad.

Sides of metazonites smooth, generally without grooves or impressions; stric-
tures sharp and distinct. Gonopores short and knob-like. Sternum of segment 4
(Fig. 1) with relatively long, apically divided process, slightly longer than widths
of adjacent coxae; sternum of segment 5 with two elevated areas between Sth
legs and distinct processes between 4th legs, coalesced medially, only slightly
shorter than widths of adjacent coxae; sternum of segment 6 with two faint knobs
between anterior legs and shallow, convex, recession between posterior legs to
accommodate gonopodal acropodites. Postgonopodal sterna with bicruciform
impressions on segments 8-10, becoming flat and plate-like posteriorly. Coxal
tubercles beginning on caudal legs of segment 9, becoming sharply pointed on
segment 10 and continuing to segment 17; prefemoral spines beginning on caudal
legs of segment 5 and continuing to ultimate leg pair, becoming progressively
longer and sharper caudally; tarsal claws hooked, not bisinuate. Hypoproct
rounded; paraprocts with margins strongly thickened.

Gonopodal aperture elliptical, 3.4 mm wide and 1.5 mm long at midpoint, in-
‘dented on anteriolateral margins, sides raised above metazonal surface. Gono-
pods in situ (Fig. 2) with acropodites crossing in midline of aperture, extending
foward slightly beyond anterior margin of aperture. Gonopod structure as follows
(Figs. 3—4): coxae moderate in size. Prefemur moderate, with moderately long,
apically blunt prefemoral process arising on anterior side, directed toward point
beyond tip of acropodite. Acropodite moderately thick and heavy, configuration
that of inverted L, tip not extending beyond level of prefemoral process; basal
zone long, about 74 of acropodite length, with row of six blunt to sharply pointed
spurs on lateral side; spurs irregularly spaced, beginning near base of basal zone
and terminating near anterior bend; anterior bend sharp and well defined, ap-
proximately a right angle; peak short, about 43 of acropodite length; medial flange
shorter than peak, margin continuous, with blunt tooth on distal corner, tooth
directed outward away from gonopod, continuous with axis of acropodite, not
projecting to level of distal extremity of peak; lateral flange subequal in width,
but slightly longer than, medial flange, extending entire length of peak, margin
wavy; solenomerite a short bisinuate process fused to undersurface of distal ex-
tremity of peak, much shorter than width of distal extremites of peak and flanges,
directed laterad, perpendicular to axis of acropodite. Prostatic groove crossing
to lateral side on proximal portion of basal zone, running along lateral side of
acropodite stem and under surface of peak to opening at distal corner of solen-
omerite.

Male paratypes.—The male paratypes agree with the holotype in somatic fea-
tures. On the gonopods the tooth on the medial flange is missing in several spec-
imens, and the prefemoral process is more globose basally in some. Otherwise,
the gonopods are very similar, and all possess six spurs on the basal zone. The
distance between spurs and their relative lengths vary slightly.

Female paratype.—Length 44.3 mm, maximum width 10.4 mm, W/L ratio 23.5%,
depth/width ratio 69.2%. Agreeing closely with holotype in somatic details except

466 PROCEEDINGS OF THE BiOLOGICAL SOCIETY OF WASHINGTON

paranota more strongly depressed, giving appearance of more highly arched dor-
sum, and caudolateral corners of paranota more rounded.

Cyphopodal aperture extending around 2nd legs, sides raised above metazonal
surface. Cyphopods in situ with corner of receptacle and valves visible in aper-
ture, valves directed caudad, oriented dorsoventrally in body. Receptacle mod-
erately large, cupped around ventral end of valves, surface rugulose. Valves
moderate in size and subequal, surface finely granulate. Operculum minute, sit-
uated under dorsal corner of valves.

Variation.—The most variable aspects of the haerens gonopod are the config-
urations of the medial flange and prefemoral process, and the number, position,
and shape of the spurs. In a few males the peak is also longer than that of the
holotype and overhangs and extends slightly beyond the level of the prefemoral
process. Variation in the configuration of the medial flange is as described under
male paratypes; the tooth is vestigial or absent from about % of the individuals
collected, and in a few specimens it is longer and more pronounced than in the
holotype. The prefemoral process is elongate in males collected near Fairview,
Buncombe County, but the configuration in the holotype is about average for the
species. The spurs range in number from six to eight and may be more or less
evenly spaced along the basal zone, or clustered near the distal extremity. They
range from sharply pointed and subspiniform to low and blunt; usually the basal
and distalmost spurs are smaller than the others. The configuration of the soleno-
merite is mostly as described for the holotype, although it is considerably nar-
rower in a few males.

Distribution.—A subrectangular area ranging from the vicinity of Old Fort,
McDowell County, and Black Mountain (town), Buncombe County, in the north,
to just south of US highway 74 in Hickorynut Gorge, Henderson County, in the
south (Fig. 14). North Carolina state highway 9, connecting Black Mountain and
Bat Cave, bisects the range. The species is clustered in the northern and southern
extremes of the range. It is relatively common near Swannanoa, Black Mountain,
and Old Fort, in the north, and in the Hickorynut Gorge area between Fairview
and Bat Cave, in the south. An apparent distributional hiatus occurs in east-
central Buncombe County. Specimens were examined as follows:

NORTH CAROLINA: McDowell Co., 4.4 mi. N Old Fort, along co. rd. 1409,
0.2 mi. W ject. co. rd. 1408, 2M, 10 September 1977 (NCSM A1724); and 4.0 mi.
N Old Fort, along US Forest Serv. Rd. in Pisgah Nat. For., M, 10 September
1977 (NCSM A1723). Buncombe Co., 3.2 mi. NW Black Mountain, along co. rd.
2476, 1.7 mi. N jct. co. rd. 2474, M, 5 September 1977 (NCSM A1702); Black
Mountain, M, September 1901, collector unknown (RLH) and 5M, 4F, date and
collector unknown (MCZ); 2 mi. N Swannanoa, along co. rd. 2427, 1.0 mi. N jet.
co. rd. 2429, M, 5 September 1977 (NCSM A1703); 5.2 mi. NE Asheville, along
co. rd. 2424, 1.0 mi. N jet. co. rd. 2419, M, 5 September 1977 (NCSM A1704); 8
mi. SE Fairview, W side Hickorynut Gap, 2M, 2 August 1973, R. L. Hoffman
and L. S. Knight (RLH); 12.2 mi. SE Asheville, along co. rd. 2806, 0.1 mi. W
jet. co. rd. 2809, M, 12 July 1976 (NCSM A989); and 12.4-12.6 mi. SE Asheville,
along US hwy. 74, 0.4—1.0 mi. W Henderson co. line 10M, 7F, 12 July 1976 and
9 September 1977 (NCSM A979, A988, and A1721) TYPE LOCALITY. Hen-
derson Co., 2 mi. NE Gerton, along co. rd. 1606, 0.2 mi. W jct. co. rd. 1605, M,
13 September 1977 (NCSM A1739); 3.7 mi. NW Bat Cave, along US hwy. 74,

VOLUME 95, NUMBER 3 467

0.8 mi. E jct. co. rd. 1598, 2M, 3F, 12 July 1976 (NCSM A1005); 1 mi. S Bat
Cave, along US hwy. 64, Hickorynut Gap, M, 2F, 13 July 1962, R. L.Hoffman
(RLH); 1 mi. SW Bat Cave, along US hwy. 64, jct. co. rd. 1514, 2M, 4F, 12 July
1976 (NCSM A1000); 9.6 mi. NE Hendersonville, along co. rd. 1569, 0.3 mi. W
jet. co. rd. 1595, M, 2F, 12 July 1976 (NCSM A999); and 6 mi. E Fletcher, along
co. rd. 1569, 1.0 mi. W jet. co. rd. 1594, M, 2F, 13 September 1977 (NCSM
NIST):

Remarks.—Prionogonus haerens occupies a much larger area than either of its
congeners. It is the only species in the northern half of the generic range, and it
extends into the southern half to south of Hickorynut Gorge. The other two
Species occur only in the southern and eastern fringes of the range, and they are
the only “‘sigmoid’’ species in this area. The area inhabited by haerens abuts that
of the next species and traverses parts of the ranges of three other sympatric
‘sigmoid’ xystodesmids: Sigmoria rubromarginata (Bollman) in McDowell and
eastern Buncombe counties, the area occupied by intergrades; S. stibarophalla
Shelley, along NC highway 9 in eastern Buncombe County; and S. stenogon
Chamberlin, in northern Henderson County. Thus, species determinations from
within the range of haerens must be done carefully on mature males only; hae-
rens, rubromarginata, and stibarophalla all have red paranota and stripes and

can easily be confused. Sigmoria stenogon, which has purple paranota and stripes
in most of its range (Shelley 1981a), takes on a reddish tint in northern Henderson
County, and it too can be confused with haerens. Although haerens occurs sym-
patrically with these three species, they are not microsympatric, and haerens has
never been collected along with another ‘‘sigmoid’’ xystodesmid at a single site.
Thus, the competitive exclusion principle, which holds for species of Sigmoria
(Shelley 1981a), also applies for species of Prionogonus.

Prionogonus divaricatus, new species
Figs. 5-8

Type-specimens.—Male holotype (NCSM A2035) and four male and three fe-
male paratypes collected by R. M. Shelley and W. B. Jones, 7 June 1978, from
Polk County, NC, 4.8 mi. NW Mill Spring, along county road 1138, 0.3 mi. W
junction of country road 1156. Male and female paratypes deposited in FSCA.

Diagnosis.—Characterized by narrow separation of solenomerite from distal
extremity of peak; tooth of medial flange generally large, projecting distad to
about distal extremity of peak; spurs large and numerous, extending around an-
terior bend to base of lateral flange on peak.

Holotype.—Length 46.2 mm, maximum width 11.7 mm, W/L ratio 25.3%, depth/
width ratio 59.0%. Segmental widths as follows:

collum 8.1 mm 10th—14th 11.4
2nd 9.1 15th lela
3rd 10.1 16th 10.4
4th 10.9 17th 8.9
Sth 11.2 18th 6.7

6th—9th 11.7

468 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 5-8. Prionogonus divaricatus: 5, Process of 4th sternum of holotype, caudal view; 6, Gon-
opods in situ, ventral view of paratype; 7, Left gonopod of holotype, medial view; 8, The same,
lateral view. Scale line for Fig. 6 = 1.00 mm; line for other Figs. = 1.00 mm for each.

Color in life: paranota red; metaterga black with wide, red, transverse stripes
along caudal edges connecting paranotal spots; collum with red stripes along both
anterior and caudal edges.

Somatic features similar to those of haerens, with following exceptions: ©

Width across genal apices 5.4 mm, interantennal isthmus 1.9 mm. Antennae
reaching back to caudal edge of third paranota, relative lengths of antennomeres
2>3>4>5=6>1>7. Facial setae as follows: epicranial and interantennal
absent, frontal 1-1, genal I—-1, clypeal about 8-8, labral about 13-13, merging
with clypeal series and continuing for short distance along genal margins, 4 setae
on each side.

Collum broad, extending well beyond margins of following tergite. Paranota
strongly depressed, continuing slope of dorsum and angling sharply ventrad, cau-

VOLUME 95, NUMBER 3 469

dolateral corners rounded through segment 5, blunt on segments 6—14, becoming
progressively more acute posteriorly.

Process of 4th sternum (Fig. 5) shorter than widths of adjacent coxae and deeply
divided apically; knobs and elevated areas on 5th sternum similar to those of
haerens, knobs betwen 4th legs more distinctly separated; 6th sternum moder-
ately recessed between 7th legs to accommodate ends of acropodites, 7th legs
set slightly farther apart than 6th. Postgonopodal sterna bicruciately impressed
on segments 8-9, becoming plate-like with shallow central impressions poste-
riorly. Tarsal claws bisinuately curved.

Gonopodal aperture elliptical, 4.2 mm wide and 2.3 mm long at midpoint, in-
dented on anteriolateral margin, sides elevated above metazonal surface. Gono-
pods in situ (Fig. 6, of paratype) with acropodites crossing at midlengths, curving
Over Opposite sides of aperture and anterior margin to between caudal legs of
segment 6. Gonopod structure as follows (Figs. 7-8): prefemoral process short
and blunt, directed toward tip of acropodite. Acropodite moderately thick and
heavy, configuration that of inverted L, peak overhanging and extending slightly
beyond level of prefemoral process; basal zone proportionately shorter than that
of haerens, about % of acropodite length, with spur basally on medial side at
juncture with prefemur and row of 10 or so sharply pointed spurs on lateral
margin; spurs irregular in length and irregularly spaced, continuing around an-
terior bend to base of lateral flange on peak, 16 spurs total; anterior bend sharp
and well defined, approximately 90°; peak long, about % of acropodite length;
medial flange arising distal to anterior bend, terminating at distal extremity of
peak, margin smoothly continuous, with sharply pointed tooth on distal corner,
tooth directed outward away from gonopod, continuous with axis of acropodite,
projecting distad to level of distal extremity of peak; lateral flange subequal in
width to medial flange, arising near midlength of peak, terminating at distal ex-
tremity, margin slightly but smoothly indented; solenomerite a long, blunt, bi-
sinuate process, length about 34 of width of distal extremity of peak, detached
and narrowly separated from peak. Prostatic groove crossing to lateral side on
proximal portion of basal zone, running along lateral side of acropodite stem and
undersuface of peak to opening at tip of solenomerite.

Male paratypes.—The male paratypes agree with the holotype in all particulars.

Female paratype.—Length 44.9 mm, maximum width 11.1 mm, W/L ratio 24.7%,
depth/width ratio 66.7%. Agreeing essentially with males in somatic details, ex-
cept paranota more strongly depressed, creating appearance of more highly arched
body.

Cyphopods in situ with corner of receptacle and valves visible in aperture,
valves directed caudad, oriented dorsoventrally in body. Receptacle large, cupped
over ventral end of valves, surface rugulose. Valves moderate and unequal, inner
one slightly larger, surface finely granulate.

Variation.—The gonopods of divaricatus are quite uniform. Males from Ruth-
erford County display a bifurcate prefemoral process, and the basal tooth on the
medial side of the basal zone is absent from all males except those in the type
series. In all specimens the spurs extend beyond the anterior bend onto the peak
of the acropodite. The number of spurs varies from about 8 to 14, and they are
located mostly distal to midlength of the basal zone. The arch of the acropodite
is constant, overhanging and extending slightly beyond the level of the prefemoral

470 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

process in all males, and the basal zone and peak are roughly equivalent in length,
each comprising approximately % of the total length of the acropodite.

Distribution.—A small area centering on the contiguous corners of Henderson,
Polk, and Rutherford counties, ranging to the eastern edge of the Blue Ridge
escarpment in Polk County. Specimens were examined as follows:

NORTH CAROLINA: Rutherford Co., Lake Lure (town), Bottomless Pool
area, 4F, 13 July 1976 (NCSM A1011) and 3M, 5F, 9 September 1978, W. B.
Jones (NCSM A2427). Henderson Co., 1.6 mi. SE Edneyville (6.1 mi. NE Hen-
dersonville), along co. rd. 1719, 1.8 mi. NE jet. co. rd. 1720, M S5F, 8 June 1978,
R. M. Shelley and W. B. Jones (NCSM A2043). Polk Co., 7.6 mi. NW Mill Spring,
along co. rd. 1163, 1.4 mi. W jet. co. rd. 1161, M, F, 14 September 1977 (NCSM
A1743); 7.2 mi. NW Mill Spring, along co. rd. 1162, 0.8 mi. W ject. co. rd. 1161,
M, 2F, 7 June 1978, R. M. Shelley and W. B. Jones (NCSM A2034); and 4.8 mi.
NW Mill Spring, along co. rd. 1138, 0.3 mi. W ject. co. rd. 1156, 5M, 3F, 7 June
1978, R. M. Shelley and W. B. Jones (NCSM A2035) TYPE LOCALITY.

Remarks.—The gonopods of divaricatus are similar in appearance to those of
haerens, and one might justifiably wonder if the two are not subspecifically re-
lated. However, I think that the differences, coupled with their contiguous ranges
in northeastern Henderson County, indicate reproductive isolation. Among the
gonopodal differences are the following: the length of the acropodal arc, which
is longer in divaricatus and overhangs the prefemoral process to a greater degree;
fusion of the solenomerite to the undersurface of the peak in haerens, whereas
they are separate in divaricatus; the more pronounced tooth on the medial flange
of divaricatus; the greater number of spurs in divaricatus and their extension
around the anterior bend onto the peak; and the longer prefemoral process of
haerens. Somatically, the paranota of divaricatus are more strongly depressed
in males, and the process of the 4th sternum is also shorter than the widths of
the adjacent coxae, whereas it is longer than the coxae in haerens. These are
considerable differences between species whose ranges are contiguous, and there
is no possibility for clinal gradation between them. This geographic pattern is
markedly different from patterns displayed by other ‘‘sigmoid’’ species that are
divided into subspecies, species such as S. /atior (Brolemann), S. rubromarginata
(Bollman), and S. nigrimontis (Chamberlin) (Shelley 198la). The evidence there-
fore supports my belief that haerens and divaricatus are reproductively isolated,
thus valid species.

Prionogonus thrinax, new species
Figs. 9-12

Type-specimens.—Male holotype (NCSM A2041) and two male and one female
paratypes collected by R. M. Shelley and W. B. Jones, 8 June 1978, from Hen-
derson County, NC, 3.1 mi. NE Flat Rock, along county road 1802, 0.7 mi. E
junction of county road 1801. One male and one female paratypes collected by
W. B. Jones, 9 September 1978, from Henderson County, along county road
1802, 2 mi. E junction of county road 1801. Male paratype deposited in FSCA.

Diagnosis.—Distinguished from congeners by trifurcate termination of acro-
podite, comprised of tooth of medial flange, subspiniform solenomerite, and a
lateral process, tooth directed subperpendicularly to axis of acropodite, others

VOLUME 95, NUMBER 3 471

directed along axis and away from gonopod; also characterized by smoothly
continuous arc of acropodite with broad, poorly defined anterior bend; reduction
in size and number of spurs on basal zone, and by reduction of lateral flange.

Holotype.—Length 40.8 mm, maximum width 9.7 mm, W/L ratio 23.8%, depth/
width ratio 61.9%. Segmental widths as follows:

collum 6.6 mm 15th 9.3

2nd 8.0 16th 8.9

3rd 9.0 17th 8.0

4th—Sth 9.4 18th 6.0
6th—14th 97

Color in life: paranota red; metaterga black with wide, red, transverse stripes
along caudal edges connecting paranotal spots; collum with red stripes along both
anterior and caudal edges.

Somatic features similar to those of haerens, with following exceptions:

Width across genal apices 4.6 mm, interantennal isthmus 1.5 mm. Antennae
reaching back to middle of 3rd paranota, relative lengths of antennomeres 2 >
3>4=5>6>1>7. Facial setae as follows: epicranial, interantennal and
frontal absent, genal 1-1, clypeal about 12-12, labral about 15-15.

Collum broad, extending slightly beyond ends of following tergite. Paranota
moderately depressed, continuing slope of dorsum, caudolateral corners rounded
through segment 6, blunt on segments 7-14, becoming progressively more acute
posteriorly.

Process of 4th sternum (Fig. 9) small, much shorter than widths of adjacent
coxae, with four long apical setae; knobs and elevated areas of 5th sternum
distinct, similar to condition in haerens; sternum of segment 6 moderately re-
cessed between 7th legs to accommodate ends of acropodite. Postgonopodal ster-
na with bicruciform impressions on segments 8-10, becoming flat and plate-like
posteriorly with shallow central impressions. Tarsal claws bisinuately curved.

Gonopodal aperture elliptical, 3.4 mm wide and 1.8 mm long at midpoint, in-
- dented on anteriolateral margin, sides flush with metazonal surface. Gonopods
in situ (Fig. 10, of paratypes) with acropodites lying generally over opposite side
of aperture, not really crossing each other and extending only short distance
beyond anterior edge of aperture. Gonopod structure as follows (Figs. 11-12):
prefemoral process short, subtriangular, directed toward tip of acropodite. Ac-
ropodite relatively thin but sturdy, well sclerotized, configuration a smoothly
continuous arc with broad diameter, not sharply divided into zones, extending
slightly beyond and overhanging prefemoral process, terminating in three widely
separated acute projections—tooth of medial flange, solenomerite, and lateral
process; basal zone of indeterminate length, with only 4 to 5 spurs on lateral
margin; spurs greatly reduced to size of small tubercles, barely detectable in
lateral view, relatively close together, located entirely on basal zone; anterior
bend broad, poorly defined, location indeterminate; peak of indeterminate length,
terminating in long, sharply pointed solenomerite and shorter, blunter, lateral
process, widely separated from solenomerite; medial flange arising distad on ac-
ropodite (peak zone?), expanding slightly proximad, with relatively long, acute

472 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

‘

Figs. 9-12. Prionogonus thrinax: 9, Process of 4th sternum of holotype, caudal view; 10, Gono-
pods in situ, ventral view of paratype; 11, Telopodite of left gonopod of holotype, medial view; 12,
The same, lateral view. Scale line for Fig. 10 = 1.00 mm; line for other Figs. = 1.00 mm for 11-12,
1.18 mm for 9.

tooth on distal corner, tooth directed downward toward coxa, subperpendicular
to axis of acropodite; lateral flange absent, perhaps represented by lateral terminal
process of acropodite, peak not noticeably expanded on lateral side; solenomerite
long and sharply pointed, subspiniform, recurved slightly on underside, directed
along main axis of acropodite; lateral terminal process shorter than solenomerite
but longer than tooth of medial flange, relatively wide basally, subacute apically,
widely separated from solenomerite. Prostatic groove crossing to lateral side on
proximal portion of basal zone, running along lateral side of acropodite stem and
undersurface of peak to opening at tip of solenomerite.

Male paratypes.—The male paratypes agree with the holotype in all particulars.

Female paratype.—Length 44.6 mm, maximum width 10.2 mm, W/L ratio 22.9%,

VOLUME 95, NUMBER 3 473

depth/width ratio 69.6%. Agreeing with holotype in most somatic features except
collum not extending beyond ends of following tergite, and paranota more strong-
ly depressed, creating appearance of more highly arched body.

Cyphopods with corner of receptacle and valves visible in aperture, valves
directed caudad, oriented dorsoventrally in body. Receptacle smaller than those
of congeners, with two lobes, one anterior and other lateral to valves, not cupped
over ventral end of valves, surface rugulose. Valves moderate in size and sub-
equal, surface finely granulate.

Variation.—The spurs on the lateral side of the basal zone are much larger and
more pronounced in other males of thrinax. One male from Henderson County
has a slight lateral expansion on the peak, indicative of a reduced lateral flange,
but all other specimens lack any indication of this lamella. The tooth on the medial
flange in the male from Polk County is greatly reduced and bifurcate apically,
and this individual also has a much longer prefemoral process.

Distribution.—Known only from a small area in eastern Henderson and western
Polk counties near the Hungry River, a tributary of the Green River. Specimens
were examined as follows:

NORTH CAROLINA: Henderson Co., 6.6 mi. ENE Hendersonville, along co.
rd. 1734, 1 mi. SW jet. co. rd. 1525, M, F, 8 June 1978, R. M. Shelley and W.
B. Jones (NCSM A2042); 3.1 mi. NE Flat Rock, along co. rd. 1802 0.7 mi. E jet.
co. rd. 1801, 3M, F, 8 June 1978, R. M. Shelley and W. B. Jones (NCSM A2041)
TYPE LOCALITY; and 4.5 mi. NE Flat Rock, along co. rd. 1802, 1.2 mi. W
Polk co. line, M, F, 9 September 1978, W. B. Jones (NCSM A2432). Polk Co..,
4.0 mi. NNE Saluda, along co. rd. 1154, 1.2 mi. E Henderson co. line, M, F, 9
September 1978, W. B. Jones (NCSM A2431).

Remarks.—The gonopods of thrinax bear a remarkable resemblance to those
of Furcillaria aequalis Shelley, which occurs in piedmont South Carolina (see
Shelley 1981b, Figs. 3-4). If the medial flange of thrinax were displaced proximad,
the two gonopods would be very close indeed. The lateral terminal process of
thrinax is therefore analogous to the tibial process of F. aequalis, both being
terminal and subparallel to the solenomerite, but I do not consider the projections
homologous. Thus, I did not name the process in thrinax, so as not to imply
homology. Moreover, the lateral terminal projection is not a generic feature of
Prionogonus, as it is of Furcillaria and Dynoria, but rather a specific trait only
of thrinax. I do not believe that this resemblance of thrinax and F. aequalis
reflects generic affinity between Prionogonus and Furcillaria; the phenotypic
similarity at the specific level seems to be a convergence. It is noteworthy that
such different evolutionary branches as the Sigmoria-Prionogonus-Hubroria and
Furcillaria-Dynoria lines can independently produce species with such similar
gonopods.

Ecology

Prionogonus is a typical montane “‘sigmoid’’ xystodesmid genus, and its species
therefore inhabit the same rhododendron cove and river-bottom environments as
most species of Sigmoria. This habitat was described in detail by Shelley (198 1a).
The millipeds are found under thin layers of leaves of hardwood species associ-
ated with rhododendron, particularly dogwood and red maple, on relatively hard
substrates near water sources. Prionogonus divaricatus, which occurs east of its

474 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

congeners on the Blue Ridge escarpment, begins to penetrate Piedmont-type en-
vironments at lower elevations. Rhododendron was scarce at a few sites where
it was taken, and the type locality in Polk County, which is near the base of the
escarpment, had very little rhododendron and seemed to exhibit more piedmont
than montane characteristics. However, divaricatus does not extend east onto
the Piedmont Plateau proper and cannot be classified as either a truly Piedmont
species or as a transprovincial one. It is really a montane xystodesmid that can
tolerate more open and drier conditions than most cove dwellers, and can there-
fore survive in marginal environments combining mountain and piedmont features
such as occur on the escarpment. The species was not found east of the type
locality, and I suspect that this site is near its eastern range limit.

The type locality of haerens was also somewhat atypical in lacking a permanent
water source. This locality is on the western side of the Tennessee Valley divide
that marks the boundary between Buncombe and Henderson counties, in a head-
water drainage area of the French Broad River. No flowing water was present,
however, and the millipeds were found just off US highway 74 under leaves in
woodland. Occurrence of the normally cove-inhabiting haerens at this locality
can probably be attributed to the site’s being cool, damp, and sheltered, and
possessing all the features normally associated with rhododendron coves except
a water source.

Distribution

The range of Prionogonus (Figs. 13-14) is the smallest of any known aphelo-
riine genus, being approximately 26 miles long and a maximum of only 13 miles
wide. Only Lyrranea in central Georgia (Hoffman 1963) has a distribution nearly
this small, but this genus is monotypic. Prionogonus thus has the smallest range
of any polytypic milliped genus. The distribution of Prionogonus is also broader
in the northern half, inhabited by haerens alone, than in the southern half, which
is occupied by all three species. In the north haerens occurs from near Swannanoa
to Old Fort, a distance of about 13 miles, but the known area of thrinax, the most
southerly species, is less than 5 miles wide. Thus, the distribution of Prionogonus
is wider north of Hickorynut Gorge and US highway 74, and tapers rapidly to
the nearly point distribution of thrinax in the south, some 4 miles north of Saluda
near the Green River. This area does not correlate with any physiographic fea-
tures, and it straddles the eastern continental divide. Prionogonus haerens occurs
on both sides of the divide—in the French Broad River basin to the west, and in
the Catawba and Broad River drainages to the east. However, divaricatus and
thrinax are limited to the Broad and Green River basins, respectively, both of
which drain into the Atlantic Ocean. Although Prionogonus is a truly montane
genus, divaricatus, as mentioned in the ecology section, has spread eastward to
the base of the Blue Ridge escarpment and the western edge of the Piedmont
Plateau Province.

Hickorynut Gorge is the center of species diversity and abundance in Prio-
nogonus. The type-species occurs in the central part of the Gorge around Gerton
and Bat Cave in Henderson County, and is also common on the western side of
the Gorge in Buncombe County. Prionogonus divaricatus occurs on the eastern
side of the Gorge around Lake Lure, Rutherford County. However, neither species

VOLUME 95, NUMBER 3 475

Figs. 13-14. 13, Range of Prionogonus in North Carolina; 14, Distribution of species of Prio-
nogonus in western North Carolina. Dashed line is approximate boundary of southern Blue Ridge
Province (Blue Ridge escarpment). Dots, haerens; triangles, divaricatus; squares, thrinax. Each
symbol marks a single collecting locality except the southeasternmost one of haerens, which repre-
sents several samples in the vicinity of Bat Cave.

(nor any other “‘sigmoid’’ species) was encountered between Bat Cave and Lake
Lure. I investigated the section around Chimney Rock, Rutherford County, sev-
eral times without finding a single apheloriine milliped, male or female. This area
has suitable habitat, and I have no explanation for the apparent absence of the
dominant southern Appalachian milliped group from this section of the Gorge.
North Carolina highway 9 bisects the northern half of the generic range in
Buncombe County, where haerens occurs, and a number of county roads provide
access from this route and US highway 74 into the surrounding mountain area.
However, the only major highway south of Hickorynut Gorge is US 64, which
passes through a corner of the range near Bat Cave. County roads are also limited
in the part of the range inhabited by divaricatus and thrinax, and only one crosses
the eastern border of Henderson County between US 74 in Hickorynut Gorge
and I-26. Prionogonus thrinax was discovered in creek bottoms along this road,
but most of the area north of it in eastern Henderson and western Polk and

476 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

haerens divaricatus thrinax

IS

Rutherford counties was inaccessible and could not be sampled. Since two known
species of restricted distribution have evolved in this part of the generic range,
it seems possible that others might occur here, especially in the border mountains
of Polk-Henderson counties.

Fig. 1S. Relationships in Prionogonus.

Relationships

Generic.— Prionogonus is a “*sigmoid’’ xystodesmid milliped genus and is most
closely related to Sigmoria. Exactly how the two relate is unknown, but the
restricted distribution of Prionogonus could mean that it is either a recent deri-
vation from ‘‘sigmoid’’ stock, or a remnant of an early fauna that is in a period
of decline. The gonopods of haerens and divaricatus bear some similarity to those
of S. stibarophalla, which is sympatric with the former, in that the solenomerite
is obscured in medial view by the medial flange in all three species. Thus, it can
only be examined in lateral perspective. The two species of Prionogonus lack a
distal zone and apical curve, and the solenomerite is either fused to, or narrowly
separated from, the undersurface of the peak. Sigmoria stibarophalla has an
apical curve and distal zone, but the former is so narrow, and the latter so short,
that both are hidden in medial view by the enlarged flange. The more distal
crossing from medial to lateral sides of the prostatic groove in S. stibarophalla,
coupled with the presence of the apical curve and distal zone and the absence of
the spurs, are the principle reasons why it is not congeneric with haerens and
divaricatus and instead belongs in Sigmoria. The simlarities in the gonopods of
haerens, divaricatus, and §. stibarophalla can best be visualized by comparing
figures 4 and 8 in this paper with figure 26 in Shelley Cee) Perhaps Prionogonus
shares a common ancestry with S. stibarophalla.

VOLUME 95, NUMBER 3 477

The position of the solenomerite beneath the peak, and the absence of a distal
zone and apical curve, also suggest the condition in Cheiropus and Stelgipus in
southern Georgia and Florida. I do not mean to suggest affinity between Prio-
nogonus and these two genera, the most southerly in the tribe Apheloriini, but
the similarity deserves mention. Both Cheiropus and Stelgipus are monotypic,
and neither has been subjected to a comprehensive treatment. Likewise, the
original descriptions of both genera and their single species (Loomis 1944) are
inadequate by modern standards and must be redone. An analysis of Cheiropus
and Stelgipus might reveal whether there is a connection between them and
Prionogonus, or whether this similarity is merely coincidental.

Specific.—The common properties of the sharp anterior bend, the expanded
distal extremity of the peak, the presence of a lateral flange, and the location of
the solenomerite beneath the peak all point to a close relationship between hae-
rens and divaricatus. They appear to be only one step removed from a common
ancestor (Fig. 15); thrinax is more divergent.

Acknowledgments

I thank Richard L. Hoffman for access to material in his private collection,
and Herbert W. Levi for loan of specimens in the MCZ collection. John E.
Cooper, NCSM, read and commented on a preliminary draft of the manuscript,
and Renaldo G. Kuhler, NCSM scientific illustrator, prepared figures 2, 6, and
10. This research was supported in part by National Science Foundation Grant
No. DEB 7702596.

Literature Cited

Hoffman, Richard L. 1963. A new diplopod genus and species from Georgia (Polydesmida: Xys-
todesmidae).—Proceedings of the Biological Society of Washington 76: 113-120.

Loomis, Harold F. 1944. Millipeds principally collected by Professor V. E. Shelford in the eastern
and southeastern states.—Psyche 51:166—177.

Shelley, Rowland M. 198la. Revision of the milliped genus Sigmoria (Polydesmida: Xystodesmi-
dae).—Memoirs of the American Entomological Society No.—33, 140 pp.

——. 198lb. A new xystodesmid milliped genus and three new species from piedmont South
Carolina (Polydesmida: Xystodesmidae).—Proceedings of the Biological Society of Washington
94:949_967.

North Carolina State Museum of Natural History, P.O. Box 27647, Raleigh,
North Carolina 27611.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 478-483

NAUSHONIA PANAMENSTS, NEW SPECIES (DECAPODA:
THALASSINIDEA: LAOMEDITDAE) FROM THE
PACIFIC COAST OF PANAMA, WITH |
NOTES ON THE GENUS

Joel W. Martin and Lawrence G. Abele

Abstract.—Naushonia panamensis is described from a Pacific Panamanian mud
flat. It is the fifth described species of the genus and the first recorded from the
tropical eastern Pacific. A revised key to the genus is presented.

The genus Naushonia Kingsley, 1897, was reviewed recently by Goy and Pro-
venzano (1979). At that time four species were known: N. crangonoides Kingsley,
1897, ranging from Woods Hole, Massachusetts, to Bogue Sound, North Caro-
lina; N. portoricensis (Rathbun, 1901) collected at Puerto Rico, Cuba, Bermuda,
and Quintana Roo, Mexico; N. perrieri (Nobili, 1904) known from French So-
maliland, Red Sea; and N. macginitiei (Glassell, 1938) known from La Jolla,
California, and Sonora, Mexico. Goy and Provenzano (1979) were able to examine
at least three specimens of each species except N. perrieri, which was unavail-
able. Apparently species in this genus are either rare or difficult to collect. The
Species are known from only a few specimens each and there are probably two
or three additional species known from larvae only (see review by Goy and
Provenzano 1979).

While at the Smithsonian Tropical Research Institute Marine Laboratory in
Panama in 1972, one of us (LGA) collected a single male specimen of an appar-
ently undescribed species of Naushonia, which we describe here. Description of
this species extends the known range of the genus into the tropical eastern Pacific.

Drawings were made with the aid of a Wild M-S stereoscope and a Wild M-11
compound stereoscope, with camera lucida. Because only a single specimen was
collected, the mouthparts are left intact. Carapace length (including rostrum),
carapace width, and total length are abbreviated CL, CW, and TL respectively.

Naushonia panamensis, new species
Figs. 1-3

Material.—Panama: Eastern Pacific, west bank of Panama Canal; in mud flat
off mangroves, about 200 m north of Interamerican Bridge of Panamanian High-
way; 4 November 1972; coll. L. G. Abele, J. B. Graham; | male, CL 6.2 mm,
CW 3.7 mm, TL 14.7 mm (Holotype, USNM 190735).

Description.—Carapace (Figs. 1, 2A, 2D) cylindrical with linea thalassinica
pronounced; anterior and posterior median dorsal carinae, separated by deep,
distinct cervical groove. Anterior dorsal carina weak. Postorbital tooth strong,
smooth, and simple. Anterolateral tooth strong and acute with small posterior
spine. Margins of carapace lightly setose. Anterior part of carapace with paired
rows of small spines roughly circumscribing gastric region and converging ante-

VOLUME 95, NUMBER 3 479

Fig. 1. Naushonia panamensis, male holotype, lateral view.

riorly onto base of rostrum. Rostrum triangular, slightly down-curved and me-
dially depressed, tipped with sparse setae and several small spines.

Eyes well developed, clearly visible from above.

Abdomen slightly longer than carapace. Pleura of segments 1-3 broadly round-
ed ventrally; pleura of segments 5 and 6 nearly straight ventrally. Pleopods absent
on first abdominal segment; other pleopods biramous, lanceolate, and with plu-
mose setae.

Telson (Fig. 2B) broadly rounded, margin with single lateral spine at about
midlength and many plumose setae. Uropods (Fig. 2B) with complete transverse
sutures, most obvious on medial branch. Lateral branch with 3-4 lateral spinules
proximal to lateral end of transverse suture; medial branch with single lateral
spine at end of suture. Both branches with many plumose marginal setae.

Antennae almost as long as body (14.0 mm), multiarticulate, lightly setose.

Antennal scale (Fig. 2C) ovate with 4 lateral and 2 terminal teeth, several lightly
plumose lateral setae and about 20 medial, long, plumose setae.

First pereopods (Figs. 1, 3E) large, slender, subchelate, and subequal. Merus
with 5 ventral and 6 dorsal teeth; carpus without teeth and with shallow distal
groove; propodus with 2 strong proximal ventral teeth, 1 midventral tooth, 1 large
and 2-3 small distal ventral teeth, and 5 small teeth located along distal % of
dorsal margin. Dactylus long, slender, heavily setose, with 6 small dorsal prox-
imal teeth.

All pereopods (Fig. 3) with many simple setae. Second pereopod shorter and
more robust than third to fifth, its dactylus thicker and more setose than those
of remaining posterior pereopods and bearing about 35 movable ventral spines.
Dactyli of pereopods 3—5 with ventral row of about 50-70 movable spines.

480 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

|

Fig. 2. Naushonia panamensis, male holotype: A, Dorsal view of carapace; B, Telson and uro-
pods, setae not illustrated; C, Antennal scale; D, Anterior of carapace, dorsal view. Scale bar = 1.0
mm for A, B, and D; bar = 0.5 mm for C.

Habitat.—The single male specimen was collected from a small puddle (ap-
proximately 10 x 20 x 3 cm) on a large exposed mud flat about 50 meters sea-
ward of a red mangrove swamp. The salinity was 24%c, temperature 28°C, al-
though the salinity in this area can vary substantially (Glynn 1972).

Remarks.—Naushonia panamensis is the fifth described species of the genus
and extends the known range of the genus to the tropical eastern Pacific; the
other Pacific Naushonia is N. macginitiei, known from southern California (Glas-
sell 1938) and Sonora, Mexico (Goy and Provenzano 1979).

The following combination of characters will distinguish N. panamensis from
other species in the genus: rostrum triangular, armed with small lateral spines;
postorbital tooth simple; telson with | lateral spine at about midlength; antennal

VOLUME 95, NUMBER 3 481

Fig. 3. Naushonia panamensis, male holotype, pereopods: A-a, Fifth pereopod and dactylus; B-
b, Fourth pereopod and dactylus; C-c, Third pereopod and dactylus; D-d, Second pereopod and
dactylus; E, First pereopod, setae not illustrated. Scale bar = 1.0 mm.

scale with 4 marginal and 2 terminal teeth; propodus of first pereopod long and
slender with length more than three times width; dactylus of first pereopod armed
with 6 teeth on proximal, superior margin; dactyli of pereopods 2—5 armed with
numerous (35-70) movable spines.

482 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

The key presented by Goy and Provenzano (1979) can be modified to include
Naushonia panamensis as follows:

1. Uropods with complete transverse sutures.
A. Linea thalassinica pronounced, carinae of the carapace weak.
a. Telson without lateral spine; antennal scale with 10 or more mar-

@inal teeth ek: ears eres ee ee eee eee eee N. crangonoides
b. Telson with lateral spine; antennal scale with less than 10 (holotype
with 6)mareinal teeth ss) ass. ele Soe eee N. panamensis
B. Linea thalassinica not pronounced, carinae of carapace well devel-
oped.

a. Telson with lateral spine; antennal scale with 6 marginal teeth, dis-
tal tooth largest and recurved inward; lateral movable spines on

dactylus of fourth pereopod only ................. N. portoricensis
b. Lateral movable spines present on dactyli of all 3 posterior pereo-
DOGS, BAILS Se 5 na ites oes eBeies.«. awelacGantine a toy che te Aenean eee N. perrieri

2. Uropods with incomplete transverse sutures; uropodal exopodites with 2
lateral spines, margin ending with 2 small spines and stout movable spine
ee ra arr es tS in Se a be EE 5 An oa ids 5 oe N. macginitiei

Although we have not examined the specimens seen by Goy and Provenzano,
completeness of the suture of the uropods seems a variable taxonomic character;
although this suture is complete in N. panamensis, it is weak and indistinct
toward the medial margin (as in the illustration of N. portoricensis by Goy and
Provenzano), especially on the lateral branch. Also, relative development of the
linea thalassinica is not clear to us from the illustrations of Goy and Provenzano
(1979:354). Perhaps a simpler key, based on the illustrations of Goy and Proven-
zano and those of Glassell (1938) and Chace (1939), and incorporating our single
specimen of N. panamensis, 1s:

1. Rostrum acute to broadly rounded.

A. Propodus of first pereopod with 2 strong proximal ventral teeth; dac-
tylus of first pereopod with proximal superior teeth; postorbital spine
SUMMPIS,, 3%. hc ad saci eee Mocha lc dso 320s Ga eRe ee N. panamensis

B. Propodus of first pereopod without 2 strong proximal ventral teeth,
with either no teeth below midventral tooth or a row of small spines;
dactylus of first pereopod without superior proximal teeth; postorbital
spine bifid or trifid.

a. Antennal scale with 10 or more marginal teeth; uropodal lateral
branchswith S*Outer Spimes .: 2.2. = seein 2 ere N. crangonoides
b. Antennal scale with fewer than 10 marginal teeth; uropodal lateral
branch with 2 outer spines. |
1. Terminal teeth of antennal scale strongly curved medially
BES ARS tet inser ya ote a eM RRP NE ou Bai N. portoricensis
2. Terminal teeth of antennal scale straight, not strongly curved
WN CANIS ee sss ce ok wn bce. ee ee eee N. macginitiei
2. Rostrum truncate, with serrate anterior margin ............... N. perrieri

Etymology.—After the type-locality.

VOLUME 95, NUMBER 3 483

Literature Cited

Chace, F. A., Jr. 1939. On the systematic status of the crustacean genera Naushonia, Homoriscus,
and Coralliocrangon.—Annals and Magazine of Natural History (11)3:524—530.

Glassell, S. A. 1938. New and obscure decapod Crustacea from the west American coasts.—
Transactions of the San Diego Society of Natural History 8(33):411-454.

Glynn, P. W. 1972. Observations on the ecology of the Caribbean and Pacific coasts of Panama.
In The Panamic biota: some observations prior to a sea-level canal, M. L. Jones, Editor.—
Bulletin of the Biological Society of Washington, no. 2, 270 pp.

Goy, J. W., and A. J. Provenzano, Jr. 1979. Juvenile morphology of the rare burrowing mud shrimp
Naushonia crangonoides Kingsley, with a review of the genus Naushonia (Decapoda: Thal-
assinidea: Laomediidae).—Proceedings of the Biological Society of Washington 92:339-359.

Department of Biological Science, Florida State University, Tallahassee, Flor-
ida 32306.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 484-494

PARALVINELLA GRASSLEI, NEW GENUS, NEW SPECIES
OF ALVINELLINAE (POLYCHAETA: AMPHARETIDAE)
FROM THE GALAPAGOS RIFT GEOTHERMAL VENTS

Daniel Desbruyeres and Lucien Laubier

Abstract.—Several unknown or newly described animals forming a flourishing
community, have been collected in the immediate vicinity of the warm hydro-
thermal springs east of the Galapagos Islands. Among them, a polychaetous an-
nelid collected within the tubemass of vestimentiferan worms represents a new
genus and species, Paralvinella grasslei, placed in the subfamily Alvinellinae,
recently erected within the family Ampharetidae for Alvinella pompejana Des-
bruyeres and Laubier, 1980, a species collected at another hydrothermal area on
the East Pacific Rise at 21°N. Paralvinella grasslei exhibits fewer morphological
and biological adaptations than A. pompejana. The two species are compared. It
is suggested that they originated from a single ancestral population that became
divided into isolated subpopulations.

One of the major biological discoveries of the last decade is the rich and unusual
fauna clustered around thermal springs at centers of rapid seafloor spreading, at
depths from 2500 to 2700 meters. The thermal springs were first discovered on
the Galapagos Rift in February 1977, using the research submersible Alvin (Bal-
lard 1977). Early in 1979, more geothermal vents were discovered west of the
original area, as well as further north along the East Pacific Rise spreading zone
near the Mexican coast at latitude 21°N. All active vents were found to be sur-
rounded by very dense populations of benthic animals, such as mussels, present
only on the Galapagos Rift, clams, and vestimentiferan tubeworms of unusually
large size (Galapagos Biology Expedition Participants 1979). 7

In a recent paper (Desbruyéres and Laubier 1980), we described a new genus
and species of tubicolous polychaete collected during the RISE cruise of the
research submersible Alvin (April-May 1979) on the East Pacific Rise at 21°N.
In spite of several aberrant features, the polychaete, named Alvinella pompejana,
was placed in the family Ampharetidae (order Terebellida) and in a new subfam-
ily, Alvinellinae, which exhibits remarkable biological features, in particular the
existence of two successive ontogenetic states.

Alvinella pompejana, very abundant at the East Pacific Rise vents, has not
been observed at the Galapagos Rift vents, although some species such as the
clam Calyptogena magnifica Boss and Turner, 1980, the crab Bythograea ther-
mydron Williams, 1980, and the vestimentiferan tubeworm Riftia pachyptila Jones,
1981, are common in both hydrothermal areas. From a biogeographical point of
view it can be emphasized that these unusual species belong basically to the same
faunal pool and derive from a single evolutionary trend.

While the description of Alvinella pompejana was in press, we received from
Dr. Frederik Grassle a single specimen of Alvinellinae found among washings
from vestimentiferan worms at the ‘‘Rose Garden’’ site, Galapagos Rift. Sur-

VOLUME 95, NUMBER 3 485

prisingly, this new species differs markedly from A. pompejana. According to
the generic features usually recognized within the Ampharetidae, these differ-
ences clearly merit the erection of a second genus within Alvinellinae. Even
though it might be considered somewhat injudicious to create a new genus and
species based on a single specimen, the biogeographical and evolutionary aspects
of this unexpected discovery are considered sufficiently important to justify a
preliminary description. While this paper was being reviewed, a second specimen
from the same station was found by Dr. Meredith L. Jones in a vestimentiferan
tube, confirming our first conclusions.

Order: Terebellida
Family: Ampharetidae Malmgren, 1865
Subfamily: Alvinellinae Desbruyéres and Laubier, 1980

Paralvinella, new genus

Type-species.—Paralvinella grasslei.

Etymology.—Derived from the generic name Alvinella which is dedicated to
the Alvin team. |

Diagnosis.—Body long and posteriorly attenuated. Prostomium extremely re-
duced, lacking appendages. Numerous smooth oral tentacles invaginable in buc-
cal cavity, together with paired ventral structures. Body with 2 indistinct regions
separated only by slight decrease in diameter; thoracic or abdominal regions not
characterized by definite structures. Four pairs of branchiae on 4 segments, first
achaetous, last 3 setigerous. Paleae absent. Setigerous segment 7 modified, with
large prominent acicular hooks. Some anterior notopodia with digitiform dorsal
lobe. Notosetae capillary and smooth when viewed with light microscope. Sessile
uncinigerous tori starting on setiger 15 and continuing posteriorly. Uncini ar-
ranged in single rows, each with only 2 unequal teeth. Pygidium rounded, without
appendages.

The main characteristic features of Paralvinella agree well with the diagnosis
of the subfamily Alvinellinae: first segment achaetous with branchiae, one ante-
rior segment modified, parapodia normally biramous, with capillary notosetae
and uncinigerous neuropodial tori. However, presence of ventral paired struc-
tures in the buccal apparatus would lead to a slight change in the subfamily
definition. It seems premature to propose an emended diagnosis at the moment.

Paralvinella grasslei, new species
Figs. 1, 2

Type-locality, material examined.—Two specimens collected during Alvin dive
990, 7 December 1979, Galapagos Rift, ‘“Rose Garden’’ area, 2451 meters depth,
00°48'25'"N, 86°13’48’”W. Holotype found among washings of vestimentiferan tubes,
second specimen recovered in empty tube of a young adult Riftia. Holotype
deposited in the collections of the Division of Worms, National Museum of Nat-
ural History, Smithsonian Institution, Washington, D.C. (USNM 67703).

Etymology.—This species is named for Dr. Frederik Grassle, Woods Hole
Oceanographic Institution, Chief Scientist of the Galapagos Biology Expedition.

Description.—Holotype 43 mm in length and 4.8 mm in greatest width (anterior

486

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Paralvinella grasslei: A, Uncinus in lateral view; B, Habitus in latero-ventral view with

oral tentacles extended; C, Anterior end in dorsal view, right branchiae removed; D, Anterior end in
ventral view, showing paired structures and oral tentacles. ;

VOLUME 95, NUMBER 3 487

third), with 99 setigerous segments. Second specimen 54 mm in length and 4 mm
wide, with 104 setigerous segments. Color after preservation in ethanol pinkish,
slightly iridescent, with normal and modified setae bright yellow. Body comprises
large anterior part, followed by decrease in diameter between setigers 47 to 52,
posterior part slender and tapering to pygidium (Fig. 1B). No ventral shields.

Prostomium extremely reduced, devoid of glandular ridges or ocular patches,
completely hidden by branchiae. Peristomium with 2 large lateral lobes enclosing
prostomium laterally, and ventral lip bearing paired lateral hollows and paired
transverse glandular crests (Fig. 1D). Buccal apparatus comprising large number
of smooth oral tentacles, each with marked ciliated groove along one side, and
two ventral paired structures, each arising from single short median stem, ending
in 3 rounded lobes (Fig. 1D). (It was not possible, without greatly damaging the
animals, to elucidate the insertion of the oral tentacles and paired structures, nor
to confirm the morphology of the prostomium.)

First segment achaetous, clearly separated from cephalic region and fused dor-
sally with next 3 setigerous segments; faint transverse lines ventrally. Next 3
setigerous segments with paradodia reduced to notopodia with capillary setae
disposed in latero-dorsal line. Branchial region formed by first 4 segments with
4 pairs of branchiae, together with dorsal part of slightly expanded fifth segment.
_Branchiae all similar, strong and regularly attenuated, stem with large number of
slender filaments irregularly inserted on 2 opposite narrow areas of stem (Figs.
1B, C; 2A, C). Branchial filaments with 2 opposed lines of small pores, 0.10 wm
in diameter with long, slender secondary filaments arising from them. Morpho-
logical detail same as in branchial lameilae of Alvinella pompejana.

Next 11 segments (segments 5 to 15, setigers 4 to 14) also with parapodia
reduced to notopodia (except the modified segment). Posterior segments (from
segment 16 or setiger 15) with notopodia and neuropodia with single rows of
uncini. Setigers 6 to 15 with dorsal digitiform lobe on notopodium increasing in
size from setiger 6 to maximum on setigers 10-13, then decreasing (Fig. 1B).
Segment 8 (setiger 7) strongly modified, lacking notopodium. On right side of
holotype, neuropodium with 3 very large acicular hooks directed posteriorly; no
hooks at all on left side but empty socket is clearly visible (probably damaged).
On left side of paratype neuropodium with 4 very large acicular hooks directed
posteriorly; no hooks visible on right side but they can be seen internally using
X-radiography. Following segment not modified. Notopodia of anterior unmodi-
fied segments well developed, provided with 10 to 15 capillary notosetae per
notopodium. Notosetae covered with continuous layer of minute spinelets when
viewed under scanning electron microscope (Fig. 2D). In cross section, notosetae
with regular network of hexagonal fibers 0.5 wm in diameter with median cylin-
drical canal.

From segment 16 (setiger 15), uncinigerous neuropodial tori present and con-
tinuing to end of body, without developing pinnules. Number of uncini per row
25—45, in single rows, with teeth facing anteriorly (retrogressive situation). Uncini
with one main tooth surmounted by smaller secondary tooth (Fig. 1A), as in A.
pompejana.

From segments 48 to 53, body diameter decreasing abruptly from 4.8 mm to
2.7 mm, with no other morphologically distinctive features. Pygidium rounded,
lacking appendages, with anus opening in center.

488 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

VOLUME 95, NUMBER 3 489

Ethology and ecology.—The 2 specimens were found without parchment-like
tubes; the holotype was found among washings of vestimentiferan tubes, the
second specimen was recovered from the empty tube of a young adult Riftia.
Both come from the same dive on the “‘Rose Garden.’’ The ‘‘Rose Garden’’
geothermal vent area is characterized by dense beds of the vestimentiferan Riftia
pachyptila Jones, 1981, living near geothermal vents from which flow warm water
at 15° to 20°C, and probably using internal symbiotic bacteria and/or their me-
tabolites as a source of organic carbon. Compared with Alvinella pompejana,
Paralvinella grasslei shows no morphological adaptations which might be related
to the uptake of dissolved organic material. From several measurements of tem-
perature at the Galapagos Rift areas, a temperature of 20°C seems to be the
highest to which P. grasslei is exposed; on the contrary, colonies of Alvinella
pompejana seem to be restricted to waters with temperatures ranging upward
from 30°C. Alvinella pompejana is probably adapted to peculiar ecological con-
ditions that do not occur at the Galapagos Rift.

Comparison of P. grasslei and A. pompejana

For the following reasons P. grasslei is comparable to the juvenile of A. pom-
pejana: they both show body constrictions at about the same distance from the
- anterior end, from segments 49 to 54 in A. pompejana, and from segments 48 to
53 in P. grasslei; they both are only slightly attenuated in the posterior regions,
in contrast to the abrupt tapering at the posterior end of the adult A. pompejana.
This conclusion does not imply the existence of a different adult form for P.
grasslei. However, there are some arguments against the existence of a compa-
rable juvenile form in P. grasslei: the posterior part of P. grasslei does not show
any modification comparable to the notopodial extensions and their ciliated tips
in the case of the juveniles of A. pompejana; also, there is nothing comparable
in P. grasslei to the necrotic dorsal area of the constricted region of A. pompe-
jana. In fact, it seems highly probable that there is only one form in P. grasslei,
and that this form is not a juvenile of the sort found in A. pompejana.

Apart from this biological comparison, which does not alone justify the erection
of a new genus, the 2 species differ as follows: there is a lobate buccal structure,
in addition to the oral tentacles, in P. grasslei; the prostomium is extremely
reduced in P. grasslei; the peristomium is well-developed, with 2 lateral expan-
sions, in P. grasslei; there are filamentous branchiae in P. grasslei, lamellate
branchiae in A. pompejana; the branchial region is composed of 4 segments, the
first achaetous and the next 3 setigerous in P. grasslei, and of one achaetous
segment only in A. pompejana; there is a single modified anterior segment with
acicular setae in P. grasslei (segment 8), while there are 2 in A. pompejana
forming a more complex system (segments 5 and 6); some anterior notopodia of
P. grasslei bear digitiform dorsal lobes while such lobes are lacking in A. pom-
pejana; the notosetae of the 2 species are of differing structure.

a

Fig. 2. Paralvinella grasslei: A, B, Detailed and general view of branchial filament; C, Isolated
branchia; D, Part of capillary notoseta. (Scanning electron micrographs)

490 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Comparison of diagnostic characters of the two species of Alvinellinae.

Prostomium

Peristomium

Buccal structures

Branchial region

Type of branchiae

Modified segments with acicu-
lar hooks

Anterior notopodia

Posterior notopodia

Dorsal part of segments

Ontogenetic stages

Notosetae

Tube

Occurrence and distribution

Paralvinella grasslei

Extremely reduced

Well-developed, with 2 lat-
eral expansions

Lobate paired structure
with oral tentacles

4 pairs of branchiae on 4
segments, first achaetous,
three setigerous

Filamentous

Segment 8 (setiger 7)

With digitiform dorsal lobes

Not modified

Not modified

One ontogenetic stage only,
corresponding to juvenile
stage of A. pompejana

Capillary, smooth when
viewed under light micro-
scope

Unknown

Among vestimentiferan
tubes, Galapagos hydro-
thermal area, temperature
10 to 20°C

Alvinella pompejana

Reduced
Reduced

Oral tentacles only

4 pairs of branchiae on first
achaetous segment

Lamellate

Segments 5 and 6 (setigers
and 4 and 5)

Digitiform notopodial lobes
lacking

Complex ciliated digitiform
notopodial extensions on
juveniles

Covered with dense tubular
structures in adults

Two different ontogenetic
stages, the juvenile stage
with twice number of seg-
ments of adult stage

Capillary, geniculate, with 2
longitudinal rows of alter-
nating teeth

Parchment-like tube cov-
ered with particles of me-
tallic sulfide

In spongy masses on hydro-
thermal vents, East Pacif-
ic Rise near 21°N, tem-
perature at least 32°C

These various characters, compared with the conventional diagnostic features
of the Ampharetidae, fully justify the erection of a new genus for the Galapagos
Rift polychaete, and serve as well to reinforce the isolated position of the subfam-
ily Alvinellinae within the family. The structure of the uncini with only 2 teeth
in the Alvinellinae is not at all typical of the Ampharetidae, and is probably one
of the more important characteristics of the aberrant Alvinellinae.

This comparison between P. grasslei and A. pompejana enables one to con-
clude that they are much closer to one another than to any other genus of Am-
pharetidae. The phylogenetic relationships between P. grasslei and A. pompejana
can be elucidated from the structure of the branchial region and the position of
the modified anterior segment. General trends of evolution in the Ampharetidae
include reduction of branchial segments by the cephalization process (Day 1964):

VOLUME 95, NUMBER 3 49]

in the primitive situation, there is one pair of branchiae on each branchial segment
(ordinarily 4 branchial segments in the Ampharetidae). This process of cephali-
zation could very well apply to P. grasslei and A. pompejana, assuming that the
4 distinct branchial segments of P. grasslei are fused to give the single achaetous
segment in A. pompejana. Cephalization usually does not affect more posterior
specific features, such as the position of modified segments.

This is clearly the case relative to P. grasslei (modified segment being segment
8) and A. pompejana (first modified segment being segment 5). This suggests the
direction of the evolutionary relationship between the species, P. grasslei being
more primitive than A. pompejana.

Taken as a whole the differences between the two species fully justify the
erection of a new genus for the Galapagos polychaete (Table 1).

Apart from the systematic and phylogenetic relationships, another comparison
must be made from a biological point of view, i.e., the feeding behavior and
corresponding morphological adaptations. Like most Ampharetidae, P. grasslei
is probably a surface deposit-feeder (Fauchald and Jumars 1979). It has oral
tentacles as well as buccal structures; the latter are probably more efficient than
the tentacles and enable the animal to catch larger particles. The situation is quite
different for A. pompejana, which, in addition to the oral tentacles, has special
tubular structures along the body of the adult; they can probably be considered
aS an adaptation for taking up released dissolved organic material (Des-
bruyeres and Laubier 1980:272). It is impossible to estimate the importance of
this uptake of dissolved organic material compared with surface deposit-feeding.
Still, A. pompejana lives very close to high temperature hydrothermal vents,
much closer than most other species of the hydrothermal community, the vesti-
mentiferan Riftia pachyptila being the other species close to the vents. Organic
material released by bacterial activity in the immediate vicinity of the vents could
be an important trophic source for A. pompejana, but there is no evidence for
this as yet. Suggestions that chemo-autotrophic bacteria present in the hydro-
thermal waters are a major food source for filter-feeding organisms such as the
mytilid mussel from the Galapagos Rift hydrothermal area have been proposed
(Rau and Hedges 1979). More recent data indicate that organic nitrogen of nu-
tritional importance for vent animals is initially synthesized within the vent en-
vironment and this synthesis may be preceded by N, fixation (Rau 1981).

Biogeographical and Evolutionary Remarks

It is obviously difficult in the present stage of our knowledge to establish the
evolutionary relationships between P. grasslei and A. pompejana: both are known
from single and limited locations, and P. grasslei by 2 specimens only. The direct
distance between the Galapagos Rift and the East Pacific Rise near 21°N, is
approximately 1800 nautical miles. Following the axis of the Rift and Rise, where
hydrothermal activity can occur, the distance is much greater, nearly 2200 nau-
tical miles. Due to the relatively large size of both species of Alvinellinae and the
recent progress in sampling deep-sea benthic organisms, there is very little doubt
that they live only in the immediate vicinity of hydrothermal vents where a supply
of organic matter is made available through chemo-autotrophy. Another pertinent
argument is given by the discovery of large accumulations of dead giant clams

492 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(Calyptogena magnifica) in areas of the East Pacific Rise, 21°N, where hydro-
thermal vents are no longer active (Francheteau et al. 1980:76-77). Those clam
cemeteries associated with ‘“‘dead’’ vent areas clearly demonstrate the basic role
of hydrothermalism as the direct or indirect energy source of these animal com-
munities (Enright er al. 1981).

Assuming that both species of Alvinellinae, as well as certain benthic organ-
isms, are strictly linked with hydrothermal vents and bacterial chemo—autotro-
phy. one has to face an apparent paradox among the ““hydrothermal community”’
species: a mollusk Calyptogena magnifica, the vestimentiferan Riftia pachyptila,
and the crab Bythograea thermydron are present in both locations; others, such
as the mytilid mussel, and the enteropneust, are restricted to the Galapagos Rift
area (perhaps this reflects insufficient knowledge of the vent community at 21°N);
the Alvinellinae form the only known example of speciation from a common
ancestor resulting in 2 different species, one for each hydrothermal site. An ex-
planation could be found in the different dispersal capabilities of the ‘“hydro-
thermal community’’ components. It has been demonstrated, in the case of the
new genus and species of Mytilidae from the Galapagos Rift, based on studies of
the larval shell morphology of juvenile specimens, that these animals have a long
planktonic larval life (Galapagos Biology Expedition Participants 1979; Lutz et
al. 1980). Abyssal currents may transport the larval stage hundreds of kilometers.
This is probably not the case with the Alvinellinae. In the family Ampharetidae,
the few existing data on reproduction and larval development show that the eggs
are usually deposited in the tube of the female, and that larval development is
non-pelagic, larval stages beyond the 3-setiger stage being found outside the mother
tube, on the mud surface. This has been observed for the Ampharetinae (Zottoli
1974). as well as for the Melinninae (Nyholm 1950).

Even the free benthic larval stage seems to be rather short, probably a few
days or a few hours. For Amphicteis floridus, it has been shown in the laboratory
that 3-setiger larvae begin tube-building just after leaving the tube of the female
(Zottoli 1974). Dispersal capabilities thus appear very limited. On the other hand,
biogeographical data show that several species of Ampharetidae, mainly from
shallow waters, have a very wide distribution, for example Amphicteis gunneri,
Anobothrus gracilis, Ampharete acutifrons, Melinna cristata. Even a truly deep-
sea species such as Amphicteis sargassoensis Hartman and Fauchald has been
found in the Atlantic Ocean from Walvis Ridge in the southeastern part to the
northwestern part. In the present state of knowledge of the Alvinellinae, it seems
that the following hypothesis can account for these two opposite sets of data:
even with their very low dispersal capabilities, the genetic flow is sufficient to
prevent rapid speciation when the physical environment constantly provides suit-
able conditions for life. The special physical environment provided by the hy-
drothermal activity is not likely to be constant: due to their strict and distinct
adaptation to this environment the two species of Alvinellinae probably originated
from a single ancestral population and were isolated over time into a series of
sub-populations where speciation occurred independently. Alvinella pompejana
and P. grasslei must be considered as two examples of this process, and one can
expect other new taxa to be found in different locations where hydrothermal vents
occur. This hypothesis leads to the following remark: at the present time, since
the intermediate morphological types that could exist between the two known

VOLUME 95, NUMBER 3 493

species of Alvinellinae have not been found, it seems practical to establish two
distinct genera, fully justified on the basis of conventional taxonomical criteria.
It must be stressed that the generic criteria serving to differentiate A. pompejana
and P. grasslei are the ones ordinarily used within the family Ampharetidae.

Résumé

A proximité immédiate des sources hydrothermales chaudes découvertes a lest
des iles Galapagos, ont été récoltés divers organismes animaux souvent inconnus
et formant une communauté exubérante. Parmi eux, une Annélide polychete a
été récoltée dans les amas de tubes de Vestimentifera. Cette Annélide appartient
a une espece nouvelle, Paralvinella grasslei gen. sp. nov., et doit étre rangée
dans la sous-famille des Alvinellinae recemment créée au sein des Ampharetidae
pour renfermer Alvinella pompejana Desbruyeres et Laubier, 1980, espece ré-
coltée sur un autre site hydrothermal de la ride du Pacifique oriental par 21°N.
Paralvinella grasslei présente des adaptations moins marquées que celles de A.
pompejana. Une comparaison entre ces deux formes est faite et une hypothese
sur leurs liens phylogénétiques est proposée.

Acknowledgments

We thank F. J. Grassle, Woods Hole Oceanographic Institution, for placing at
our disposal the first specimen sorted from a sample studied by H. Sanders and
Nancy Copley, M. H. Pettibone, Smithsonian Institution, for reviewing this
manuscript, and M. L. Jones for sending us the second specimen and reviewing
the paper. This is Contribution n° 25 from the Galapagos Rift Biology Expedition,
supported by the National Science Foundation and Contribution n° 755 from the
Centre Océanologique de Bretagne.

Literature Cited

Ballard, R. D. 1977. Notes on a major oceanographic find.—Oceanus 20(3):35—44.

Boss, K. J., and R. D. Turner. 1980. The giant white clam from the Galapagos Rift, Calyptogena
magnifica species novum.—Malacologica 20:161—194.

Day, J. H. 1964. A review of the family Ampharetidae (Polychaeta).—Annals of the South African
Museum 48:97-120.

Desbruyéres, D., and L. Laubier. 1980. Alvinella pompejana gen. sp. nov. Ampharetidae aberrant
des sources hydrothermales de la ride Est-Pacifique.—Oceanologica Acta 3:267—274.

Enright, J. T., W. A. Newman, R. R. Hessler, and J. A. McGowan. 1981. Deep-ocean hydrothermal
vent communities.—Nature 289:219-221.

Fauchald, R., and P. A. Jumars. 1979. The diet of worms: a study of polychaete feeding guilds.—
Oceanography and Marine Biology Annual Review 17:193—284.

Francheteau, J., T. Juteau, D. Needham, and C. Rangin. 1980. Naisance d’un océan.—CNEXO
édition, Strasbourg 1980; 88 pp.

Galapagos Biology Expedition Participants. 1979. Galapagos 79: Initial findings of a deep-sea bio-
logical quest.—Oceanus 22(2):2-10.

Jones, M. L. 1981. Riftia pachyptila, new genus, new species, the vestimentiferan worm from the
Galapagos Rift geothermal vent (Pogonophora).—Proceedings of the Biological Society of
Washington 93:1295-1313.

Lutz, R., M. Jablonski, D. Rhoads, and R. Turner. 1980. Larval dispersal of a deep-sea hydro-
thermal vent bivalve from the Galapagos Rift.—Marine Biology 57:127-133.

Nyholm, K. 1950. Contribution to the life-history of the ampharetid, Melinna cristata.—Zoologiska
Bidrag fran Uppsala 29:79-91.

494 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Rau, G. H. 1981. Low 1°N/!4N in hydrothermal vent animals: ecological implications.—Nature 289:

484-485.

, and J. I. Hedges. 1979. Carbon-13 depletion in a hydrothermal vent mussel: suggestion of

a chemosynthetic food source.—Science 203:648-649.

Williams, A. B. 1980. A new crab family from the vicinity of submarine thermal vents on the
Galapagos Rift (Crustacea: Decapoda: Brachyura).—Proceedings of the Biological Society of
Washington 93:443—472.

Zottoli, R. A. 1974. Reproduction and larval development of the ampharetid polychaete Amphicteis
floridus.—Transactions of the American Microscopical Society 93:78-79.

Centre Océanologique de Bretagne, B.P. 337, 29273 Brest Cédex, France.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 495-504

A NEW GENUS OF BOMOLOCHID COPEPODS FROM
INDO-WEST PACIFIC NEMIPTERID FISHES

Roger F. Cressey

Abstract.—A new genus, Holocolax, is described to include H. nemipterus
(Pillai, 1973) and two new species, H. longisetus, and H. russelli. The new genus
is characterized by an armature of | spine and 3 long setae on the last endopod
segment of the female third leg as opposed to 2 spines and 2 setae in other
bomolochid genera. The parasites are described from 3 species of Nemipterus.

In 1973 Pillai described a new species of bomolochid copepod from the inner
surface of the operculum of Nemipterus japonicus (Bloch) from India. Following
the revisions of bomolochid genera by Vervoort (1962, 1969) he assigned the new
species to the genus Holobomolochus Vervoort on the basis of the first antenna
being without modified setae. He noted, however, that the new species differed
from all other species of Holobomolochus by the peculiar nature of the last
-endopod segment of the female third leg. In other species of Holobomolochus
the last endopod segment bears 2 short, inner spines and 2 setae. Pillai’s species
has 1 short inner spine and 3 setae. Examination of a number of species of
Nemipterus resulted in recovering 2 more species, closely related to H. nemipteri
with the same peculiar armature of the female third leg.

Based on these additional 2 new species described below I consider the 3
species to represent a new genus.

Holocolax, new genus

Diagnosis.—Bomolochidae. Body form typical of family. Thoracic segments
bearing legs 2—5 free. Abdomen of female 3-segmented; male 2-segmented. Caudal
rami of female with 5 minor and 1 major setae; male with 4 minor and 2 major
setae. Rostrum without hooks. First antenna 5-segmented. Second antenna with
3 terminal and | subterminal claw, all of about equal length. Mouthparts typical
of family. Maxilliped of female without accessory process. Legs 1-4 biramose,
all rami 3-segmented except male leg 4 endopod 2-segmented. First leg of female
modified as typical of family, male unmodified. Middle endopod segment with 1
inner seta and last endopod segment of leg 3 with 1 short spine and 3 setae in leg
3 of female, male with 2 spines and 2 setae.

Etymology.—A combination of Holobomolochus, the genus to which Pillai’s
species was assigned, and colax, a common suffix in bomolochid genera. Gender
masculine.

Type-species.—Holocolax longisetus, new species.

Holocolax longisetus, new species
Figs. 1-21

Material examined.—Holotype 2 (USNM 190510), allotype ¢ (USNM 190511)
and 22 paratypes 2 (USNM 190512) from the gill area of 26 Nemipterus mulloides

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

496

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VOLUME 95, NUMBER 3 497

(USNM 200270) collected by Kunz and Wells from Taiwan. Additional material
consisting of 5 specimens from 7 specimens of the same host species (USNM
76624) from Taiwan.

Female.—Body form as in Fig. 1. Total lengths and greatest widths of 3 spec-
imens 914 x 507 um, 884 x 464 um, and 812 x 449 um (widths measured at
widest part of cephalon, additional measurements based on largest specimen).
Genital segment (Fig. 2) wider than long (144 x 86 wm). Abdomen 3-segmented,
segments measure 58 X 85 wm, 58 X 86 um, and 44 xX 82 um (I X Ww) respec-
tively; last abdominal segment with 2 large ventral patches of spatulate spines
(Fig. 3). Caudal rami (Fig. 3) longer than wide (36 x 20 um), each with a ventral
patch of spatulate spinules, an outer lateral seta, an outer subterminal seta (middle
one much longer and wider at its base than other 2), and | dorsal subterminal
seta (all setae naked).

First antenna (Fig. 4) with 5 segments, 15 plumose setae on outer edge of first
2 segments, no modified setae on first segment; an aesthete on each of last 2
segments. Rostral hooks absent. Second antenna (Fig. 5) second segment with
numerous hooklike spinules arranged in irregular rows and a subterminal artic-
ulated spine about equal in length and width to the 3 articulated spines of last
segment; 2 naked setae on last segment in addition to spines. Mouthparts (Fig.
6) typically bomolochid except first maxilla with the smaller, naked seta, more
prominent than in other bomolochids. Maxilliped (Fig. 7) with a small naked seta
on first segment, 3 long plumose setae on second segment, and a heavily scler-
otized recurved claw, without an accessory process.

Legs 1-4 biramous. Leg | (Fig. 8) exopod with heavily sclerotized spine on
outer distal corner of first segment, otherwise typically bomolochid; endopod first
2 segments each with a patch of spatulate spinules along outer distal margin.
Inner seta of basipod modified as a bilobed spatulate process. Second leg (Fig.
9) basipod with prominent plumose seta at outer distal corner; exopod first seg-
ment with large patch of hairs on outer distal surface and a fringed spine at outer
distal corner, second segment with outer fringed spine and an inner seta, last
segment with 3 outer fringed spines and 6 setae, all setae on last segment with
short plumosities along outer edge, all spines of exopod with terminal flagellum;
endopod first segment with patch of spatulate spinules along outer margin and an
inner seta, second segment similar to first except an additional seta, last segment
with 2 outer plumose spines and 3 plumose setae. Leg 3 (Fig. 10) exopod first
segment with large patch of spatulate spinules on outer distal surface and a spine
at outer distal corner, second segment similar to first except with an inner seta,
last segment with patch of spatulate spinules, 3 outer spines, and 5 inner setae,
all exopod spines with fringe along outer margin and with a terminal flagellum,
setae of last segment armed as those of leg 2; endopod segments all with outer
patch of spatulate spinules, first 2 segments each with an inner seta, last segment
with 1 outer spine and 3 terminal setae (outermost longest and spinose in distal
half). Leg 4 (Fig. 11) similar to leg 3 except exopod last segment with only 4

—

Figs. 1-6. Holocolax longisetus 2: 1, Dorsal; 2, Genital segment and abdomen, dorsal; 3, Last
abdominal segment and caudal rami, ventral; 4, First antenna; 5, Second antenna; 6, Mandible,
paragnath, first and second maxillae.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

498

Figs. 7-10. Holocolax longisetus 2: 7, Maxilliped; 8, Leg 1; 9, Leg 2; 10, Leg 3.

VOLUME 95, NUMBER 3

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dorsal; 14, Last abdominal segment and caudal rami, ventral; 15, First antenna; 16, Maxilliped.

499

500 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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Figs. 17-19. Holocolax longisetus d: 17, Les 1: 18, Leg 2:19, Leg 3.

VOLUME 95, NUMBER 3 501

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AA

Figs. 20-25. Holocolax longisetus 3: 20, Leg 4; 21, Leg 5. Holocolax russelli °: 22, Leg 2; 23,

Leg 5. Holocolax nemipteri 2: 24, Leg 2 exopod; 25, Last abdominal segment and caudal rami,
ventral.

502 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

setae, endopod without hairs on outer edges of segments and last segment with
2 spines and one seta. Leg 5 (Fig. 12) free segment with patch of spatulate spinules
on outer surface and 4 setae of about equal length, terminalmost sparsely plu-
mose, others finely plumose. Leg 6 represented by 2 short setae at area of egg
sac attachment (Fig. 2).

Male.—Total length 464 um and greatest width 188 um (measured at widest
part of cephalon). Genital segment (Fig. 13) somewhat wider than long. Abdomen
2-segmented, last segment with ventral patches of long, pointed spinules (Fig.
13). Caudal ramus (Fig. 14) about as long as wide (11 um) with a ventral patch
of pointed spinules and 6 setae, terminal 2 nearly equal in width at base.

First antenna (Fig. 15) as in female except outer setae of first 2 segments not
as robust and more sparsely plumose, inner plumose seta of third segment ex-
tending well beyond tip of antenna (corresponding seta in female very short).
Other cephalic appendages as in female except maxilliped (Fig. 16) basal segment
robust with a spinose knob on inner margin, claw relatively straight with inner
edge of tip spinose opposing knob on basal segment.

Leg | (Fig. 17) not modified as in female, coxopod with row of pointed spinules
along outer distal corner. Basipod with a prominent plumose seta on outer corner,
an inner naked seta on distal margin inner to endopod, and 2 large patches of
pointed spinules; exopod first segment with patch of spinules along outer edge
and a spine on outer distal corner, second segment similar to first except with an
inner seta, last segment bearing 2 outer spines and 5 setae, outermost seta with
spinose outer edge, all spines with subterminal flagellum; endopod first and sec-
ond segments each with a patch of spinules and an inner seta, last segment with
1 outer spine and 5 setae. Leg 2 (Fig. 18) similar to leg | except basipod without
ornamentation, exopod last segment with 5 setae, endopod segments with a row
(rather than patch) of spinules on outer distal border of each, second segment
with 2 inner setae, last segment with 2 spines and 3 setae. Leg 3 (Fig. 19) similar
to leg 2 except exopod second segment lacking outer spine, endopod second
segment with one inner seta (2 in leg 2) and last segment with 2 setae (3 in leg
2). Leg 4 (Fig. 20) similar to leg 3 except exopod last segment with 4 setae (5 in
leg 3), endopod 2 segmented and last segment with one terminal seta flanked by
a spine on each side. Leg 5 (Fig. 21) with a patch of pointed spinules at inner
distal corner and 2 terminal setae, inner seta with short close plumosities, outer
sparsely plumose. Leg 6 absent.

Etymology.—ongisetus, alluding to the unusually long seta on the last seg-
ment of the endopod of the female third leg.

Remarks.—This species differs from Holocolax nemipteri (Pillai) by its patches
of scalelike spinules on the abdomen and caudal rami whereas nemipteri has
pointed spinules. It differs from both nemipteri and russelli as its terminal setae
of leg 5 are all about equal in length whereas in the other species the terminalmost
seta is much longer.

Holocolax russelli, new species
Figs. 22—23

Material examined.—Holotype 2 (USNM 190513) and | 2 paratype (USNM
190514) from the gill area of 1 specimen of Nemipterus metopias (BPBM 22159)

VOLUME 95, NUMBER 3 503

from Damaguete City, Philippines. Additional material consisting of 2 2 from 1
specimen of the same host species (USNM 227079) from Palawan, Philippines
and | 2 from the same host (CSIRO AS5/80/57) from Australia.

Female.—Total length and greatest width 798 x 431 um respectively. This new
species closely resembles H. longisetus and H. nemipteri, and only those points
of difference will be discussed. Genital segment 94 x 135 um (1 X w respec-
tively). Abdominal segments measure 59 X 88 wm, 35 X 77 um, and 41 X 65 um
(1 X w), respectively. Caudal ramus somewhat longer than wide (29 x 21 um).

Leg 2 (Fig. 22) exopod first segment with large patch of scalelike spinules on
distal half (hairs in H. longisetus and H. nemipteri) otherwise similar to H. lon-
gisetus. Leg 5 (Fig. 23) mid-terminal seta much longer than others (all setae of
equal length in H. longisetus).

Male.—Unknown.

Etymology.—This species is named for Dr. Barry Russell who identified and
sorted collections of Nemipterus species for me to examine for parasitic cope-
pods.

Remarks.—This new species can be easily separated from H. nemipteri and
H. longisetus by the presence of a large patch of spatulate spinules on the first
exopod segment of the second leg in H. russelli (hairs in H. nemipteri and H.
longisetus).

Holocolax nemipteri (Pillai, 1973)
Figs. 24-25

Holobomolochus nemipteri Pillai, 1973:487.

Material examined.—2 @ from the gill area of 6 Nemipterus japonicus and 2
2 from 5 specimens of the same host species from the Andaman Sea, R. V.
Anton Bruun, Cruise 1, Stations 28C and 49. Three 2 from 2 specimens of the
same host from Palk Bay, Sri Lanka.

Female.—This species was well described by Pillai 1973, and I will only sup-
plement that description with those characters which differ from the previous 2
Species.

Leg 2 exopod (Fig. 24) first segment with a row of hairs rather than the patch
of hair present in H. longisetus; the middle outer spine on the last segment is
longer than the ones to either side, in the other 2 species these spines are of equal
lengths. The ventral surface of the last abdominal segment and caudal rami (Fig.
25) bear patches of small spinules unlike the prominent spatulate spinules on the
abdomen and caudal rami of the other 2 species.

Acknowledgments
I thank Dr. Barry Russell for identifying and sorting host species for me and
Hillary Boyle Cressey for reading and commenting on the manuscript.
Literature Cited

Pillai, N. Krishna. 1973. Three new bomolochid parasites on fishes of the Kerala coast.—Indian
Journal of Fisheries 20(2):487-496.
Vervoort, W. 1962. A review of the genera and species of the Bomolochidae (Crustacea, Copepoda),

504 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

including the description of some old and new species.—Zoologische Verhandelingen 56:1-—
ate

1969. Caribbean Bomolochidae (Copepoda: Cyclopoida).—Studies of the Fauna of Curacao
and other Caribbean Islands 28:1—125.

Department of Invertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 505-508

OPHIACANTHA ABYSSA, NEW SPECIES, AND
OPHIOPHTHALMUS DISPLASIA (CLARK), A SUGGESTED
NEW COMBINATION IN THE OPHIUROID FAMILY
OPHIACANTHIDAE (ECHINODERMATA:
OPHIUROIDEA) FROM OFF
OREGON, U.S.A.

Michael A. Kyte

Abstract.—Ophiacantha abyssa, anew species of ophiacanthid ophiuroid from
abyssal waters off Oregon is described. It is distinguished by a proliferation of
oral papillae and aboral arm plates and oral shields bearing spines. Ophiacantha
diplasia (Clark, 1911) is placed in the ophiacanthid genus Ophiophthalmus.

An extensive benthic faunal sampling program by Dr. A. G. Carey, Jr. of
Oregon State University has produced a large collection of echinoderms. Material
from the abyssal regions included 32 specimens of an undescribed ophiacanthid
ophiuroid. These collections also contained 43 specimens of Ophiacantha dipla-
sia. Using Carey’s material and Clark’s (1911) type-specimen, it was found that
O. diplasia properly belongs in the genus Ophiophthalmus Matsumoto, 1917.

Suborder Laemophiurina Matsumoto, 1915
Family Ophiacanthidae Perrier, 1891
Ophiacantha abyssa, new species
Figs. 1, 2

Material examined.—Oregon State University, School of Oceanography sam-
ples from Tufts Abyssal Plain: OTB 104, 40°12.7’N, 126°30.3’W, 4260 m, 14
January 1966, 9 specimens; OTB 163, 44°43.2'N, 134°44.6’W, 3860 m, 1 March
1967, 2 specimens; OTB 164, 44°37.4'N, 133°39.8'W, 3699 m, 4 March 1967, 2
specimens; OTB 331, 44°40.9'N, 133°31.9'W, 3717 m, 4 June 1970, 5 specimens;
OTB 334, 44°31.1'N, 134°43.8’W, 3858 m, 5 June 1970, holotype (USNM E273 16),
2 paratypes (USNM E27317); BMT 228, 44°45.7'N, 134°23.8’W, 3354 m, 20
May 1970, 7 specimens; BMT 230, 44°25.8'N, 132°13.4'W, 3655 m, 1 June 1970,
2 specimens (all material except types is in author’s possession or at Oregon State
University).

Etymology.—abyssos (Greek)—bottomless, referring to the abyssal occurrence
of the species.

Description.—Disc diameter 18 mm; arm length approximately 80 mm. Disc
circular, covered with skin bearing thorny stumps. Radial shields completely
covered with thorny stumps, conspicuous as raised areas. Oral interbrachial spaces
nearly covered with thorny stumps as on disc. Genital slits large, conspicuous.
Oral shields pentagonal, wider than long, center concave, distal edge bearing a
varying number of thorny stumps or short spines as on disc. Adoral plates curved,
inconspicuous. Oral papillae highly variable, in one to several rows on each side
of jaw.

506 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Ophiacantha abyssa: Aboral view.

Aboral arm plates obtusely triangular with distal edges rounded; significantly
broader than long. Basal plates well separated. First 3 or more plates bear thorny
stumps on distal edges. All ventral arm plates with small tubercles or a swelling
in center. First ventral arm plates wider than long with rounded distal edges and
truncate proximal sides. Lateral arm plates large, meeting above and below,
separating aboral and oral arm plates. Each side arm plate with 8 smooth or
slightly serrate stout spines. Tentacle spines single, basal scales broad at base

VOLUME 95, NUMBER 3 507

Fig. 2. Ophiacantha abyssa: Oral view.

with pointed tip, distal scales spiniform. Color in life rust-red with black-appear-
ing oral papillae.
Distribution.—Northeast Pacific abyssal regions of the Gorda Ridge and Tufts
Abyssal Plain off Oregon and northern California, 3354 to 4260 m.
Discussion.—Ophiacantha abyssa is distinguished from most other members
of the genus by the proliferation of oral papillae and the occurrence of spines on
the distal edges of the aboral arm plates and the oral shields. Ophiacantha spec-

508 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tablis Sars and O. anomala Sars resemble O. abyssa in possessing a proliferation
of oral papillae on the oral and adoral plates. However, thorny spines do not
occur on the aboral arm plates of O. spectablis or on the oral shields of O.
anomala. Several additional differences exist including the smoothness of the
arm spines, the shape and smooth texture of the tentacle scales, and the differing
shape of aboral arm plates in O. anomala and O. spectablis. In addition, the
distribution of these two species is restricted to the eastern Atlantic above 2000
m (D’yakonov 1967, Mortensen 1933), while O. abyssa occurs in the Northeast
Pacific below 3300 m.

Ophiophthalmus diplasia (Clark), new combination
Ophiacantha diplasia Clark 1911:209.

Discussion.—After examining Clark’s (1911) type (USNM 25647) and 43 spec-
imens from a wide range of locations off Oregon, it is suggested that Ophiacantha
diplasia be placed in the genus Ophiophthalmus. This species possesses short
and rounded radial shields that have the distal one third exposed in contrast to
the elongate rectangular shields that are completely covered with skin typical in
Ophiacantha. The disc is covered with definite flat scales bearing sparse coarse
spherical granules instead of skin bearing thorny stumps or spines as in Ophia-
cantha. The large, flat tentacle scales and long, smooth glassy arm spines are
also characteristic of Ophiophthalmus rather than Ophiacantha. Arm spines in
Ophiacantha typically approximate dorsally on proximal arm segments. Arm
spines do not approximate in any of the 43 specimens from Oregon or in Clark’s
type-specimen.

Acknowledgments

I am grateful to Dr. David Pawson, Curator, Division of Echinoderms, U.S.
National Museum of Natural History, for his helpful review of an earlier version
of this paper, and to Dr. A. G. Carey, Jr. for his support and patience during the
preparation of this article. Partial support was generously provided by the U.S.
Atomic Energy Commission (contract AT (45-1) 2227 Task Agreement no. 12).

Literature Cited

Clark, H. L. 1911. North Pacific ophiurans in the collection of the United States National Mu-
seum.—Bulletin of the United States National Museum 75:1—295.

D’yakonov, A. M. 1954. Ophiuroids of the USSR seas.—Keys to the fauna of the USSR, Zoological
Institute of the Academy of Sciences of the USSR, no. 55; [translated from Russian Bi the
Israel Program for Scientific Translations, Jerusalem] 1967; 123 pp.

Mortensen, T. 1933. Ophiuroidea.—The Danish Ingolf Expedition 4(8): 1-121.

School of Oceanography, Oregon State University, Corvallis, Oregon 97331.
Present address.—527 212th Street S.W., Bothell, Washington 98011.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 509-514

MESONERILLA PROSPERA, A NEW ARCHIANNELID FROM
MARINE CAVES IN BERMUDA

Wolfgang Sterrer and Thomas M. Iliffe

Abstract.—A new species of Archiannelida, Mesonerilla prospera, is described
from inland marine caves of Bermuda. This new species is particularly interesting
in that it was collected from thin layers of mud on and under stones in totally
dark areas of the caves, whereas all other marine Archiannelida (except one deep-
water species) are members of the interstitial sand fauna.

Recent studies of island marine caves in the Western Atlantic have revealed
the presence of rich marine faunas. Comprehensive cave faunal surveys are now
underway in Bermuda (Sket and Iliffe 1980) and on San Salvador Island, Bahamas
(Carpenter 1981). Additionally, spot collections have been made from a number
of sites (Peck 1973). Significant marine troglobites discovered include Remipedia,
a new class of Crustacea from Grand Bahama Island (Yager 1981), a variety of
caridean shrimps from diverse localities (Hobbs et al. 1977), and a mysid from
Jamaica (Bowman 1976). Further collections from anchialine pools, as yet un-
classified ecologically, have produced representatives of the amphipod suborder
Ingolfiellidea (Stock 1977a), hadziid amphipods (Stock 1977b), an amphipod of the
genus Psammogammarus (Stock 1980), the isopod family Microparasellidae (Stock
1977c), and the order Thermosbaenacea (Stock 1976).

The Bermuda Islands, site of the current survey, consist of Pleistocene and
Recent eolian and marine limestones completely capping a mid-ocean volcanic
seamount. The islands’ limestone caves were primarily formed during low stands
of sea level corresponding to periods of Pleistocene glaciation (Bretz 1960, Palmer
et al. 1977, Iliffe 1981). As postglacial sea levels rose, much of the former extent
of the caves was flooded by sea water. Approximately 200 inland caves are known
from Bermuda, over half of which contain tidal, sea level pools. Bermuda’s long-
est cave is the 1.8 km, totally underwater Green Bay Cave system (Iliffe 1980).
A biological survey of the terrestrial and marine caves of Bermuda was begun in
1978 (Sket and Iliffe 1980). Although most of the animals collected from marine
caves were more or less regular immigrants from open littoral habitats, a number
of new species including blind and probably subterranean ones were found. Two
new species of caridean shrimp, Somersiella sterreri and Typhlatya iliffei (Hart
and Manning, 1981); an isopod, Atlantasellus cavernicolus, representing a new
family (Sket, 1979); and a new calanoid copepod, Miostephos leamingtonensis
(Yeatman, 1980) have so far been described from Bermuda’s caves. We here
describe a new species of Archiannelida collected from the Walsingham Caves,
Bermuda.

Mesonerilla prospera, new species
Fig. 1

Material.—Walsingham Caves, Hamilton Parish, Bermuda: 8 December 1978,
Walsingham Sink Cave, 1 specimen from 1 m water depth collected with long-

510 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Mesonerilla prospera: a, Young female specimen, dorsal view; b, Detail of setae from
segments 2-9; c, Detail of setae from segment 1; d, Anterior end, dorsal view; e, Spermatid; f, Sperm.

handled dip net; 5 January 1979, Walsingham Cave, 3 specimens from 1 m water
depth collected with long-handled dip net; 22 May 1979, Walsingham Cave, 3
specimens collected from 1 m depth with long-handled dip net; 24 July 1979,
Walsingham Cave, 3 specimens collected from 1 m depth with slurp gun, 3 spec-
imens collected from 1 m depth with small dip net; 9 October 1981, Cripplegate
Cave, 1 specimen collected from surface waters of outflowing tidal spring with
plankton net; 28 November 1981, Deep Blue Cave, | specimen collected from
12-15 m depth by hand net with scuba; 17 January 1982, Emerald Sink Cave, 1
specimen collected from 11 m depth by hand net with scuba; 26 January 1982,
Cherry Pit Cave, 4 juvenile specimens collected by pumping water from 2.5 m
depth through a plankton net; 31 January 1982, Myrtle Bank Cave, 1 specimen
collected from rock walls and ledges at 5 m depth by hand net with scuba; 23
June 1982, Cherry Pit Cave, 6 specimens collected from 1-5 m depths by hand
net. Observations and measurements from 9 specimens were used for the species
description. A whole mount of a female specimen collected on 23 June 1982
at Cherry Pit Cave has been deposited as holotype (USNM 73764).

Distribution.—Known only from the anchialine habitats of the Walsingham
Caves, Bermuda.

Habitat.—The Walsingham Caves are located in a 450 m wide strip of land
separating the nearly enclosed Harrington Sound from the more open Castle
Harbour. At least 40 cave entrances are known from the Walsingham tract, a
low, hilly, heavily overgrown area of 0.15 km?. Caves of this area are character-
ized by fissure entrances and large collapse chambers. Two large underwater
caves with a total of 1 km explored length, 20 m depths and 7 known entrances
probably represent only segments of a much larger and more complex cave sys-
tem. Walsingham and Deep Blue Caves have already been connected by cave

VOLUME 95, NUMBER 3 511

divers, as have Cripplegate and Myrtle Bank Caves. Both of these groups of
caves as well as Cherry Pit and Walsingham Sink Caves are believed to be hy-
drologically connected as part of the same system. Emerald Sink Cave, located
only 300 m away from Cherry Pit Cave, may also be connected.

Amplitude and phase of the tides have been measured in Harrington Sound,
Castle Harbour and pools of the Walsingham Caves. While there is no significant
difference between tides in Castle Harbour and those of the open sea, Harrington
Sound tides have an average lag time of 2 hours and 45 minutes and a range of
only 30% of the open sea tides. Tides for the Walsingham Caves are intermediate,
having an average | hour lag and 60% range. This difference in phase between
Harrington Sound and Castle Harbour tides produces alternating tidal currents
flowing through the caves such that an estimated 50% of the tidal volume of
Harrington Sound (half a million m*) passes through caves (Morris et al. 1977).
Plankton and organic detritus carried by tidal currents probably provide the pri-
mary source of food for the cave animals. Surface salinities in pools of the Wal-
singham Caves average 27%o, while at 1 m salinities already reach 34%o.

In order to determine where exactly within the cave pools Mesonerilla prospera
is living, selective collecting was carried out. Using a slurp gun, material was
collected from the steep rock walls of the cave. A small dip net was used to
obtain animals from the upper few centimeters of the thick silt on the cave floor.
Material from piles of 30-60 cm diameter rocks with only thin layers of silt was
collected both with the slurp gun and by moving rocks quickly up and down to
generate a flushing action and then sweeping a small net through the disturbed
water. Six specimens of Mesonerilla prospera were collected from the piles of
rock, three each with the slurp gun and net, while none were obtained from either
the cave walls or floor. The discovery of an archiannelid living in inland caves,
on and under stones, is exceptional since all other marine members of this group,
except one deep-water species (Sterrer 1968), are members of interstitial sand
fauna. The collection of four juveniles from open water in Cherry Pit Cave and
one adult from the Cripplegate tidal spring indicates that subterranean water
currents may be significant in determining the distribution of Mesonerilla pros-
pera.

Description.—The length of adults (excluding appendages) ranges from 1500 to
2050 wm, with the maximum width of 250-420 um (to 520 wm including parapodia)
somewhat behind midbody (Fig. la). The largest juvenile (i.e., without visible
gonads) was 1300 um; smaller juveniles collected in water pumped from above
the bottom measured 300 um (with 5 segments), 400 um (7 segments) and 450
wm (8 segments).

The prostomium (Fig. 1d) is rounded and carries a pair of reddish eyes and 3
tentacles dorsally, and a pair of palps ventrally. The median tentacle can be up
to 650 um long, the lateral ones to 600 wm; in most specimens however, they are
much shorter. Palps are up to 230 um long and 50-60 um wide. They are gently
curved, and are broadest near the base, tapering gradually towards the tip.

There are 9 setigerous segments. Whereas each parapodium carries 2 thin and
sometimes rather long (to 550 um) cirri in segments 2-9, the buccal parapodia
carry only 1 usually very short cirrus (30 um). A pair of anal cirri (urites) were
seen in only one of the specimens; they were short (120 um, and 30 um), and
are probably easily lost. Each parapodium carries about 20 setae arranged in 2
bundles. Setae are compound on all segments; they are of fairly equal length

512 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(shaft 170 wm, end piece 45 um) throughout segments 2-9, but longer (shaft 220
yum, end piece 90 um) on the first segment. The setae of segment | also differ in
having a pointed shaft spur whereas it is blunt in the setae of all other segments
(Fig. 1b, c).

The mouth opening is longitudinally slit-shaped and sometimes appears trian-
gular. The internal epithelia of pharynx, esophagus and rectum are ciliated. None
of the food particles present in the gut could be identified.

Sexes are separate. A female may have up to 8 mature eggs (to 200 um di-
ameter) arranged on both sides of the gut posterior to segment 4. None of the
females observed carried eggs or embryos externally. Of the two males, one had
segments 4—9 full of sperm (Fig. 1f) and spermatids (Fig. le). Sperm is about 22
yum long, with a bullet-shaped head of 2 um, and a tail of about 20 um.

The unrestrained animal glides slowly over the substratum by means of its
ciliation. Like many other Nerillidae, this species shows a rapid escape reaction
when disturbed.

Etymology.—The name of the species refers to its well-fed appearance, but is
also an allusion to Prospero’s Cave in Shakespeare’s ‘‘The Tempest,’ a play
allegedly inspired by the shipwreck of Bermuda’s first settlers.

Discussion.—The number of segments (9) and the compound setae clearly place
the new species in the genus Mesonerilla Remane, 1949. Of 7 species presently
ascribed to this genus [M. minuta Swedmark, 1959 has been assigned to another
genus (Jouin 1971)], 3 are distinct from M. prospera in being hermaphroditic; a
fourth, M. luederitzi Remane, 1949, is known only as juveniles. One of the re-
maining 3 species, M. biantennata Jouin, 1963, differs from M. prospera in lack-
ing a median tentacle. Another, M. intermedia Wilke, 1953, is characterized by
brood protection devices (‘‘elytres’’) in the female (Jouin 1968) which M. pros-
pera does not have. The most recently described species, M. ecuadoriensis
Schmidt and Westheide, 1977, from shallow sandy bottoms in the Galapagos
Islands, is also the one that most closely resembles M. prospera; there are dif-
ferences, however, in the shape of the palps, the number of parapodial cirri, the
presence of eyes, and the body size. At 2000 um, M. prospera is by far the largest
of all Mesonerilla species; of the described species M. intermedia reaches 1200
ym, most of the others only measure 1000 um or less.

Finally, the habitat in which M. prospera occurs, sets it apart from all other
marine Nerillidae, which live interstitially in sand, with the exception of Para-
nerilla limicola Jouin and Swedmark, 1965, which lives on deep mud bottoms
(Sterrer 1968). The only freshwater nerillid, Troglochaetus beranecki Delachaux,
1921, although not closely related morphologically, shares with M. prospera the
troglobitic way of existence (Pennak 1971). A biogeographic observation of in-
terest is that M. prospera has its closest relative on another oceanic island,
Galapagos, just as Bermuda’s cave decapod shrimps are most closely related to
species from Caribbean islands (Antigua, Cuba, Caymans, Bahamas), Ascension
Island and, again, Galapagos (Hart and Manning 1981). Whether this indicates
that oceanic islands, and island cave habitats in particular, are refuges for the
descendants of formerly widely distributed but now extinct marine faunas must
await further evidence.

VOLUME 95, NUMBER 3 ais

Acknowledgments

This study was supported by a National Science Foundation Grant (DEB-
8001836) to Thomas M. Iliffe and Wolfgang Sterrer. Additional support for cave
collections was provided by EARTHWATCH and the Center for Field Research,
and for cave diving by grants from the Explorers Club and the National Speleologi-
cal Society. Cave diving equipment and techniques used to conduct this study met
standards of the National Speleological Society. This paper is Contribution No.
900 from the Bermuda Biological Station for Research, Inc.

Literature Cited

Bowman, T. E. 1976. Stygiomysis major, a new troglobitic mysid from Jamaica, and extension of
the range of S. holthuisi to Puerto Rico (Crustacea: Mysidacea: Stygiomysidae).—International
Journal of Speleology 8(4):365-373.

Bretz, J. H. 1960. Bermuda: A partially drowned, late mature, Pleistocene karst.—Bulletin of the
Geological Society of America 71:1729-1754.

Carpenter, J. H. 1981. Ecology and taxonomy of marine cave invertebrates in the Bahama Islands.—
Proceedings of the Eighth International Congress of Speleology, Bowling Green, Kentucky,
USA, pp. 24-25.

Hart, C. W., Jr., and R. B. Manning. 1981. The cavernicolous caridean shrimps of Bermuda (AI-
pheidae, Hippolytidae, and Atyidae).—Journal of Crustacean Biology 1(3):441—456.

Hobbs, H. H., Jr., H. H. Hobbs III, and M. A. Daniel. 1977. A review of the troglobitic decapod
crustaceans of the Americas.—Smithsonian Contributions to Zoology 244: 1-183.

lliffe, T. M. 1980. Mid-ocean cave diving.—Underwater Speleology 7(4):46—48.

1981. The submarine caves of Bermuda.—Proceedings of the Eighth International Congress

of Speleology, Bowling Green, Kentucky, USA, pp. 161-163.

Jouin, C. 1968. Sexualité et biologie de la reproduction chez Mesonerilla Remane et Meganerilla
Boaden (Archiannelides Nerillidae).—Cahiers de Biologie Marine 9:31—52.

——. 1971. Status of the knowledge of the systematics and ecology of Archiannelida.—Smith-
sonian Contributions to Zoology 76:47—56.

Morris, B., J. Barnes, F. Brown, and J. Markham. 1977. The Bermuda marine environment.—
Special Publication Number 15, Bermuda Biological Station, St. George’s West, Bermuda,
120 pp.

Palmer, A. N., M. V. Palmer, and J. M. Queen. 1977. Geology and origin of the caves of Bermuda.—
Proceedings of the Seventh International Congress of Speleology, Sheffield, U.K. pp. 336-339.

Peck, S. B. 1973. Recent studies of the invertebrate fauna and ecology of sub-tropical and tropical
American caves.—Proceedings of the Sixth International Congress of Speleology, Olomouc,
Czechoslovakia 5:185—194.

Pennak, R. W. 1971. A fresh-water archiannelid from the Colorado Rocky Mountains.—Transac-
tions of the American Microscopical Society 90(3):372—375.

Sket, B. 1979. Atlantasellus cavernicolus n. gen., n. sp. (Isopoda Asellota, Atlantasellidae n. fam.)

from Bermuda.—Bioloski Vestnik (Ljubljana) 27(2): 175-183.

, and T. M. Iliffe. 1980. Cave fauna of Bermuda.—Internationale Revue der gesamten Hy-

drobiologie 85(6):87 1-882.

Sterrer, W. 1968. Paranerilla limicola Jouin & Swedmark (Archiannelida) von der norwegischen
und adriatischen Ktiste.—Sarsia 36:65-68.

Stock, J. H. 1976. A new genus and two new species of the crustacean order Thermosbaenacea

from the West Indies.—Bijdragen tot de Dierkunde 46:47-70.

1977a. The zoogeography of the crustacean suborder Ingolfiellidea with descriptions of new

West Indian taxa.—Studies on the Fauna of Curacao and other Caribbean Islands 55:131—-146.

. 1977b. The taxonomy and zoogeography of the hadziid Amphipoda with emphasis on the

West Indian taxa.—Studies on the Fauna of Curacao and other Caribbean Islands 55:1—130.

514 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

——. 1977c. Microparasellidae (Isopoda, Asellota) from Bonaire.—Studies on the Fauna of Cu-
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——. 1980. Amsterdam expeditions to the West Indian Islands: 8. A new cave amphipod (Crus-
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386.

Yager, J. 1981. Remipedia, a new class of Crustacea from a marine cave in the Bahamas.—Journal
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Yeatman, H. C. 1980. Miostephos leamingtonensis, a new species of copepod from Bermuda.—
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Bermuda Biological Station, Ferry Reach 1-15, Bermuda.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 515-521

TWO NEW SPECIES OF CERATONERETS (POLYCHAETA:
NEREIDIDAE) FROM ESTUARINE AREAS OF
NEW SOUTH WALES, AUSTRALIA

P. A. Hutchings and C. J. Glasby

Abstract.—Two new species of Ceratonereis, C. limnetica and C. turveyi, are
described from estuarine areas in New South Wales. Ceratonereis limnetica oc-
curs in an upstream site where freshwater conditions predominate. The two new
species belong to a group of Ceratonereis characterized by simple neurosetal
falcigers. This group appears to have radiated within Australia.

While examining material from ecological surveys carried out in estuarine areas
of southern Australia it became apparent that several species of Ceratonereis
have been mistakenly identified as C. erythraeensis Fauvel. Three of these species
were described by Hutchings and Turvey (in press) and in this paper an additional
two species are described.
The following abbreviations are used: Allan Hancock Foundation (AHF), Aus-
tralian Museum (AMW), British Museum (Natural History) (BMNH), Muséum
National d’ Histoire Naturelle, Paris (MNH), and Smithsonian Institution, Wash-
ington, D.C. (USNM).

Ceratonereis limnetica, new species
Fig. la—e

Holotype.—AMW 18668, 83 setigers, 64 mm length, 3.0 mm width. Para-
types.—2 specs., AHF Poly 1367; 2, BMNH ZB 1982.33-34; 2, USNM 071768;
2, AMW 18670; 41, AMW 18669; size range of paratypes 47 setigers, 15 mm
length, 2.0 mm width to 84 setigers, 72 mm length, 3.5 mm width. All material
from Lower Portland, Hawkesbury River, N.S.W., Australia (33°27’S, 150°54’E),
coll. Glasby, 20 Nov 81, intertidal sandy beach.

Other material examined.—Ceratonereis vaipekae Gibbs: Holotype BMNH ZB
1972.1, Aitutaki, Cook Islands stn A6 coll. Gibbs 2.9.1969, muddy sand, MTL-
LWN. Ceratonereis erythraeensis: Sénafir, island south of Suez Canal, Tadjoura
Bay (Red Sea), Tuléar, Madagascar; and Luta (=Dairen), China from MNH,
identified by Fauvel but not part of type-series.

Description.—Body flattened, of uniform width anteriorly, gradually tapering
posterior to middle setigers. Color in alcohol cream with brown granular pig-
mentation anterodorsally and prominent dorsal blood vessel. Prostomium width
approximately equal to length, with deep medial groove extending from near tip
to between anterior pair of eyes (Fig. la). Two pairs of small black eyes, anterior
pair slightly further apart. Palps with globose style and dorsal groove across base,
extending to level of frontal antennae. Four pairs of tentacular cirri, inner pos-
terior pair largest, extending to setiger 4, appearing shallowly annulated. Ever-
sible pharynx with slender, transparent brown jaws with 8 (left) and 6 (right)
exposed teeth. Paragnaths present only in maxillary ring, as brown cones ar-

516 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ee
we ae ee

Fig. 1. Ceratonereis limnetica. a, Anterior dorsal view; b, Anterior view parapodium 10; c, An-
terior view parapodium 40; d, Anterior view posterior parapodium; e, Giant simple falciger from 40th
parapodium. Scale in mm.

ranged as follows: I = 1; II = 20 in oblique band; III = 32 in broad transverse
band; IV = 26 in Y-shaped band.

Parapodial lobes bluntly conical anteriorly, more pointed and compressed pos-
teriorly. First 2 parapodia uniramous, lacking dorsal notopodial lobe; subsequent-
ly notopodial lobes of comparable size except in posterior setigers where dorsal
notopodial lobe is reduced, absent in last 2 setigers. Ventral neuropodial lobe
well developed anteriorly, extending almost as far as notopodial lobe (Fig. 1b)

VOLUME 95, NUMBER 3 S17)

reduced to a small tubercle by about setiger 25 (Fig. Ic), but increasing in size
relative to other lobes posteriorly (Fig. 1d). Dorsal neuropodial lobe with well
developed bluntly conical post-setal lobe in first 10 setigers only, similar in size
to ventral notopodial lobe but ligulate rather than conical in middle and posterior
setigers. Dorsal cirri on all setigers, 74-1 times length of dorsal notopodial lobe
in anterior and middle setigers, posteriorly elongating to 3—4 times length of dorsal
notopodial lobe before latter decreases. Ventral cirri extending %4—%2 way to tip
of ventral neuropodial lobe throughout.

Acicula black, brown at extremities. Numbers of setae in 10th, mid and pos-
terior setigers respectively as follows: notosetae 7, 5, 2 homogomph spinigers;
neurosetae dorsally 8, 5, 2 homogomph spinigers, 3, 0, 0 heterogomph falcigers
and 0, 2, | giant simple falcigers; neurosetae ventrally 9, 7, 5 heterogomph spi-
nigers and 7, 1, 0 heterogomph falcigers. Giant simple falcigers formed by an-
kylosis and fusion of teeth of heterogomph falcigers in dorsal neuropodial fascicle
over about setigers 25-30. Dorsal neuropodial falcigers in this region with inter-
mediate characteristics. Fully formed giant simple falcigers light brown, 3—4 times
diameter of other setae, bluntly hooked with a distinct tendon (Fig. le).

Anal cirri ventrally produced, extending over last 5 setigers.

Variation.—Variations not described for the holotype include prostomium length
0.9-1.1 times width. Palps extend to just short of or well past antennae. Longest
pair of tentacular cirri vary in length, extending to setigers 2-5. Jaws with 3-10
teeth. Paragnaths arranged as follows: I = 1-5; Il = 15-30 in an oblique band,
widest medially, III = 26-43 in a square or rectangular transverse band; IV =
20-43 in a Y- or V-shaped band. Relative lengths of parapodial lobes similar
throughout. Postsetal neuropodial lobes indistinct in small specimens, but oth-
erwise well developed in first 7-13 setigers. Numbers of setae in 10th, mid and
posterior setigers respectively vary as follows: notosetae 5-10, 3-8, 1-4 homo-
gomph spinigers; neurosetae dorsally 6-12, 4-9, 1-4 homogomph spinigers, 2-4,
0-1, 0 heterogomph falcigers and 0, 1-2, 1-2 giant simple falcigers; neurosetae
ventrally S—15, 2-11, 3-6 heterogomph spinigers and 2-7, 1-10, 0-1 heterogomph
falcigers. Occasionally heterogomph spinigers found in dorsal neuropodial fas-
cicle. Heterogomph falcigers in dorsal neuropodial fascicle anteriorly, developing
into giant simple falcigers by middle setigers. Fully formed giant simple falcigers
highly variable in shape, sometimes with 1-2 robust teeth surmounting main fang
occasionally with 3-4 minute teeth proximal to main fang, but most often smooth
(Fig. le). Variation in the nature of the giant simple falcigers occurs within
and between specimens and appears to be independent of the size or position
along the body of an individual.

Several gravid females were examined in the paratype material (AMW18669)
and no signs of epitokal modifications were observed.

Discussion.—Ceratonereis limnetica belongs to the small group of Ceratonereis
characterized by the presence of simple neuropodial falcigers. It may be distin-
guished from other members of this group occurring in Australia, for which de-
scriptions have recently been made (Hutchings and Turvey, in press), in having
post-setal neuropodial lobes in anterior setigers, relatively well developed ventral
neuropodial lobes in posterior setigers and in the paragnath arrangement (Table

Ih).

518 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Comparison of selected features between Australian Ceratonereis species characterized
by the presence of simple neuropodial falcigers. Those species recently described by Hutchings and
Turvey (in press) are designated by numbers.

Post-setal Posterior ventral
Simple neuropodial neuropodial neuropodial
Species falcigers lobe lobe Paragnath type and arrangement
C. limnetica variable present well devel- brown cones; I = 1-5;
(smooth- oped II = 15-30 in oblique
denticulate) band; III = 26—43 in

transverse band;
IV = 20-43 in V- or
Y-shaped band

Gena spau denticulate present absent transparent-dark brown
cones, variable in
size, paragnaths ab-
sent in Area III

C. n. sp. 2 denticulate present absent transparent rounded
cones and large elon-
gate reddish cones;
generally fewer par-
agnaths than for C.
limnetica

Crneisp.s denticulate absent small tubercle brown cones; similar
numbers and arrange-
ment to C. limnetica

C. turveyi denticulate absent small tubercle brown cones and domes
proximally by setiger I = 0-5, II = 5-15,
30 Il = 1-7, lV = 6-15

Ceratonereis erythraeensis Fauvel, 1918 from Madagascar and C. vaipekae
Gibbs, 1972, from the Cook Islands differ from C. limnetica in the paragnath
pattern and in the absence of post-setal neuropodial lobes in anterior setigers.

A large amount of Ceratonereis material from New South Wales estuaries and
lagoons has been examined and it appears that C. limnetica has a very limited
distribution, occurring only in the upper reaches of the Hawkesbury River, New
South Wales. It represents the numerically dominant species of polychaete in the
freshwater zone with densities of up to 1000 individuals per square meter, but
also occurs in lesser numbers further downstream where salinities reach 20%bo.
Salinities at the collection site may reach 8%o during periods of drought combined
with the effects of a spring tide (A. Jones, pers. comm.). Juveniles of 20 setigers
(about 2 mm length) are present throughout the year, but appear to be most nu-
merous from January to May. Smaller individuals were not sampled for.

The closely related species, C. n. sp. 3, also occurs in the Hawkesbury River,
although further seaward than C. limnetica. Unlike the latter, C. n. sp. 3 1s
common in estuaries and lagoons throughout southern Australia. The occurrence
of two morphologically similar species in the one estuary is unusual, but probably
occurs in other estuaries of Australia and supports the idea suggested by Hutch-

VOLUME 95, NUMBER 3 SN)

ings and Turvey (in press) that the group of Ceratonereis characterized by the
presence of simple neuropodial falcigers has radiated within southern Australia.
Etymology.—The name limnetica refers to the estuarine/freshwater habitat in
which this species lives.
Australian distribution.—Upper Hawkesbury River, New South Wales.
Habitat.—Fresh and brackish water, often in sandy sediment or in mud. Sub-
tidal and intertidal. Salinity range 0-—20%bo.

Ceratonereis turveyi, new species
Fig. 2a—f

Holotype.—AMW 14981, 114 setigers, 16 mm length, 1.2 mm width. Para-
types.—1 spec. AHF Poly 1368; 1, BMNH ZB1982.35; 1, USNM 071769, 3 AMW
18667; 1, 11276, range from 80 setigers 10 mm length, 0.9 mm width to 32 setigers,
4 mm length and 0.6 mm width (incomplete specimen). All material collected
from Posidonia seagrass beds, Merimbula, N.S.W., Australia (36°55’S, 149°55’E)
during 1976 by New South Wales State Fisheries.

Description.—Body flattened, widest over anterior half, gradually tapering pos-
teriorly. Color in alcohol, yellow-white with light brown pigmentation antero-
dorsally, especially on anterior prostomium, on palps, and around eyes. Prosto-
mium length about equal to width, with a shallow medial groove extending from
near tip to level with base of palps. Two pairs of reddish eyes, anterior pair
slightly further apart. Palps with large globose style and stout cylindrical base,
extending to just short of frontal antennae. Four pairs of tentacular cirri appear
deeply annulated, inner posterior pair largest, extending to setiger 4. Eversible
pharynx with slender, transparent brown jaws carrying 2 exposed teeth in addition
to long curved main fang. Paragnaths present in maxillary ring as small light
brown cones and larger dark brown cones with light brown bases, arranged as
follows: I = 5; I! = 14 in 2 rows on right and triangular patch on left; III = 7 in
transverse line; IV = 15 in V-shaped band; oral ring bare.

Parapodial lobes conical (Fig. 2a) becoming compressed and more pointed pos-
teriorly (Fig. 2b, c). Notopodial lobes and ventral neuropodial lobe more elongate
than dorsal neuropodial lobe throughout. First 2 parapodia uniramous, lacking
dorsal notopodial lobe. Dorsal notopodial lobe initially reduced, increasing pos-
teriorly to approximately same size as ventral notopodial lobe by middle setigers,
thereafter decreasing to ¥% size of ventral notopodial lobe, absent in last 15 se-
tigers. Dorsal neuropodial lobe without post-setal lobe, slightly smaller and less
elongate than notopodial lobes in anterior and middle setigers, about 4% size of
ventral notopodial lobe in posterior setigers. Ventral neuropodial lobe comparable
in size to dorsal neuropodial lobe anteriorly, reduced to small tubercle from about
setiger 30 onwards. Dorsal cirri on all setigers, 1-112 times length of dorsal no-
topodial lobe in anterior and middle setigers, posteriorly elongating to about 3
times length of dorsal notopodial lobe, before latter decreases. Ventral cirri ex-
tending '/2—-*/; way to tip of ventral neuropodial lobe throughout.

Acicula dark brown-black, hyaline at extremities. Numbers of setae in 10th,
mid and posterior setigers respectively as follows: notosetae 9, 8, 3 homogomph
spinigers; neurosetae dorsally 8, 4, 3 homogomph spinigers, 3, 0, 0 heterogomph
falcigers and 0, 1, 1 giant simple falcigers; neurosetae ventrally 4, 1, 2 hetero-

520 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

>

0-13

Fig. 2. Ceratonereis turveyi. a, Parapodium 11, posterior view; b, Parapodium 35, anterior view;
c, Posterior parapodium anterior view; d, Dorsal neuropodial falciger, parapodium 21, showing partial
fusion of blade to shaft; e, Ventral neuropodial heterogomph falciger, parapodium 21; f, Fully formed
giant simple falciger, posterior parapodium. Scale in mm.

gomph spinigers and 6, 10, 2 heterogomph falcigers. Giant simple falcigers formed
by ankylosis and rearrangement of teeth of heterogomph falcigers in dorsal neu-
ropodial fascicle over about setigers 20-25. Dorsal neuropodial falcigers in this
region with intermediate characteristics (Fig. 2d). Fully formed giant simple fal-
cigers about 2 times diameter of other setae, with an indistinct tendon and 3 small
denticles proximal to bluntly conical main tooth (Fig. 2f). Heterogomph falcigers
with very short blades in middle and posterior setigers (Fig. 2e), otherwise setae
normal. Anal cirri ventrally produced, extending over last 10 setigers.
Variation.—Variations not described for the holotype include prostomium length

VOLUME 95, NUMBER 3 521

equal to 0.9-1.2 times width. Longest pair of tentacular cirri varying in length,
extending from setigers 3-7; appearing only shallowly annulate in some small
specimens. Jaws with 2—5 teeth. Paragnaths variable in shape and color within
and between individuals ranging from light brown to dark brown cones; dark
brown paragnaths generally larger. Paragnaths arranged as follows: I = 0.5; II =
5—15 in an oblique or triangular patch; III] = 1-7 in a transverse line or irregularly
arranged; IV = 6-15 ina V-shaped or curved band. Relative lengths of parapodial
lobes similar throughout in paratype material except for dorsal notopodial lobe
which may be absent from setigers 3 and 4 in smaller specimens; dorsal notopodial
lobe absent posteriorly in last 12-15 setigers. Numbers of setae in 10th, mid and
posterior setigers respectively as follows: notosetae 5-9, 6-8, 3-4 homogomph
Spinigers; neurosetae dorsally 3-8, 2-5, 2-3 homogomph spinigers, 2-3, 0, 0
heterogomph falcigers and 0, 1, 1 giant simple falcigers; neurosetae ventrally 3-
4, 1-3, 2-4 heterogomph spinigers and 3-6, 3-10, 2-3 heterogomph falcigers.
Heterogomph falcigers in dorsal neuropodial fascicle anteriorly, developing into
giant simple falcigers by middle setigers. Anal cirri long, ventrally produced,
extending over last 7-10 setigers.

None of the material has obvious coelomic gametes.

Discussion.—In Table 1, the characteristics separating Ceratonereis turveyi
from other Australian Ceratonereis species possessing simple falcigers are shown.
Ceratonereis turveyi differs from these species in that the main fang is smooth,
the few teeth present occur proximally to the main fang (Fig. 2f).

Etymology.—This species is named for S. Paul Turvey who drew our attention
to it.

Australian distribution.—New South Wales (Merimbula).

Habitat.—Posidonia seagrass beds. The species was collected during a quan-
titative survey of the beds, the results of which are described by Collett et al.
(in press), where the species is erroneously referred to as C. erythraeensis Fauvel.
Sediment: muddy sand in predominantly marine conditions.

Acknowledgments

We thank Dr. D. George, British Museum (Natural History) and Mme. Renaud-
Mornant, Muséum National d’ Histoire Naturelle, Paris for loan of material, and
Dr. A. Jones, Ms. A. Murray, and Ms. C. Watson-Russell for providing us with
specimens and ecological data for one of the species.

Literature Cited

Collett, L. C., P. A. Hutchings, P. J. Gibbs, and A. J. Collins. (in press.) A comparative study of
the macrobenthic fauna of Posidonia australis seagrass meadows in New South Wales, Aus-
tralia.—Aquatic Botany.

Fauvel, P. 1918. Annélide polychéte nouvelle de |’ Afrique orientale.—Bulletin du Muséum d’ His-
toire Naturelle, Paris 24:503—509.

Gibbs, P. E. 1972. Polychaete annelids from the Cook Islands.—Journal of Zoology 168:199—220.

Hutchings, P. A., and S. P. Turvey. (in press.) The Nereididae of southern Australia. Transactions
of the Royal Society of South Australia.

Department of Marine Invertebrates (Worms), The Australian Museum, 6-8
College Street, Sydney, New South Wales 2000, Australia.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 522-529

ANOPSILANA CRENATA, A NEW TROGLOBITIC
CIROLANID ISOPOD FROM GRAND CAYMAN
ISLAND, CARIBBEAN SEA

Thomas E. Bowman and Richard Franz

Abstract.—Anopsilana crenata, a new blind unpigmented cirolanid isopod, is
described from a freshwater pool in West Bay Cave, a small cave at the NW end
of Grand Cayman Island. The cave is formed within limestone of the Late Pleis-
tocene Ironshore Formation, whose age is probably about 130,000 years BP. The
possibility that the 5 species of Anopsilana evolved by convergence rather than
from a common ancestor is discussed.

The genus Anopsilana (Cirolanidae) was established by Paulian and Delamare
Deboutteville (1956) for a new blind unpigmented isopod from fresh water in
Mitoho, a cave in southern Madagascar. This species, A. poissoni, is still known
only from the type-locality. Vandel (1964) expressed doubts about the validity of
Anopsilana, and Monod (1976) illustrated A. poissoni in detail from 2 specimens
given to him by Delamare Deboutteville and placed it in Cirolana. Monod ana-
lyzed point by point the statements in the original diagnosis of Anopsilana and
found nothing that excluded A. poissoni from Cirolana.

When Bruce (1981) distributed species of the cumbersome genus Cirolana among
6 genera (including 3 new genera), he recognized Anopsilana as valid because it
differed from Cirolana in having non-setigerous endopods on pleopods 3-5 (in
Cirolana the endopods of pleopods 3 and 4 are setigerous). Monod (1976) did not
include the pleopods in his analysis, presumably because they were not mentioned
in Paulian and Delamare Deboutteville’s original diagnosis of Anopsilana. Bruce
(1981) included in Anopsilana the type-species, A. poissoni, and 3 species that
he removed from Cirolana: A. luciae (Barnard 1940), A. pustulosa (Hale 1925),
and A. willeyi (Stebbing 1904). The new species described below brings the num-
ber of known species to 5.

Anopsilana crenata, new species
Figs. 1-2

Material.—Grand Cayman I., Cayman Is. (Caribbean Sea, NW of Jamaica),
West Bay Cave (known locally as ‘‘Blue Hole’’), a sink located on a limestone
rise at NW end of island (Fig. 3) 200-300 yds (180—275 m) from the sea at an
elevation of 20 ft (6.1 m) above sea level, 0.5 km NNE of green turtle farm
(Mariculture, Ltd.); leg. M. Langworthy and G. Morgan, 18 April 1980: 3 ho-
lotype (USNM 190712) and 37 32 paratypes (USNM 190713).

Etymology.—The specific name, from the Latin “‘crenatus’’ (notched), refers
to the scalloped appearance of the palm of the propus of pereopod 1.

Description.—Blind, unpigmented. Length of largest specimen 6.2 mm. Body
slightly more than 2 as long as wide, greatest width at pereonite 5. Head with
weak lateral angles, anterior margin produced into small rostrum reflexed ven-

VOLUME 95, NUMBER 3

i ()) \
i Ly at A
iy ) i) ) ih 1 iN | i |

Fig. 1. Anopsilana crenata: a, Habitus, dorsal; b, Habitus, lateral; c, Apex of telson; d,
Anterior region of head, ventral; e, Antenna | peduncle; f, Antenna 2 peduncle; g, Left mandible; h, i,
Incisors of right and left mandibles; j, k, Laciniae of left and right mandibles; 1, Maxilla 1; m, Maxilla
2; n, Maxilliped; o, Endite of maxilliped; p, Pereopod 1; q, Pereopod 2; r, Pereopod 3.

S23

524 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Anopsilana crenata: a. Pereopod 4: b. Pereopod 6: c. Pereopod 7: d. Pleopod 1: e. Pleopod
2: f. Endopod of ¢ pleopod 2: g. Apex of appendix masculina: h. Penes: i. Pleopod 3: j. Pleopod
k. Pleopod 5: 1. Uropod. ventral.

=

VOLUME 95, NUMBER 3 525

=a * CAVE
From Side

Ee West Bay Cave

vy
‘Turtle Farm

miles | . 5
bee RE ees Bes Eee |

Fig. 3. Below, Outline map of Grand Cayman Island: Upper left, NW end of Grand Cayman
Island, showing location of West Bay Cave: Upper nght, West Bay Cave in profile and from above,
from field sketches.

trally between bases of antennae | and meeting frontal lamina. Frontal lamina
pentagonal, about 1.6Xx longer than wide, not produced ventrally. posterior mar-
gin overlapped by clypeus. Clypeus short, about 0.66X length of labrum.

Pereonite | the longest: pereonites 2—5 gradually increasing in length, pereonite
6 subequal to or slightly shorter than pereonite 5, pereonite 7 distinctly shorter.
Posterior corners of coxae 2—3 rounded, of coxae 4—6 squared, of coxa 7 angular.
overlapping pleonites 1—2; all coxae with oblique carinae.

Pleonites subequal in length: pleonite | at least partly overlapped by pereonite
7; pleonite 5 overlapped laterally by pleonite 4: epimera of pleonites 2—3 pointed,
of pleonite 4 rounded. Telson triangular, about 0.75xX as long as wide, posterior
margin rounded, encompassing about 0.2 telson, armed with 8 short slender spines:
between 2 lateral spines are 2 plumose setae, between other spines are 3 plumose
setae.

526 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Antenna | nearly reaching posterior margin of pereonite 1; peduncle segments
1-2 fused dorsally, free ventrally; flagellum of 8-10 segments, last 5 segments
with 2-2-2-1-0 esthetes (counting distally). Antenna 2 reaching midlength to pos-
terior margin of pereonite 5; peduncle segment 5 only slightly longer than segment
4; flagellum with 18—22 segments.

Incisors of mandible 3-cuspate, cusps more deeply separated in right mandible;
right and left laciniae with 9 and 11 spines; molar with 18 and 21 spines on right
and left mandible, respectively. Exopod of maxilla 1 with 12 spines (1 not shown
in Fig. 11); endopod with 3 circumplumose spines. Maxilla 2 with 5 and 11 setae
on palp and exopod, respectively; endopod with 16 setae of varying lengths (only
14 shown in Fig. 11). Maxilliped with 2 retinacula.

Pereopod 1, palm of propus armed with low rounded scales giving it a scalloped
appearance. Pereopods 2-7 slender, successively longer, ischium, merus, and
carpus with distal groups of strong spines.

Penes low, rounded, slightly wider than long, close together.

Pleopod 1 endopod with slightly concave lateral margin. Pleopod 2 endopod,
appendix masculina bow-shaped, tapering to narrowly rounded apex, extending
beyond endopod by nearly 0.2 its length.

Uropods reaching slightly beyond apex of telson; exopod slightly shorter and
about 0.6 as wide as endopod, lateral margin with 5 spines, medial margin with
3 spines; apical notch asymmetrical with longer medial side. Endopod triangular,
lateral margin with 2 spines, medial margin with 4 spines; apical notch nearly
symmetrical, with slightly longer medial side.

Comparisons.—Of the known species of Anopsilana, A. luciae, A. pustulosa,
and A. willeyi differ from A. crenata in having well developed eyes and in living
in estuarine or marine habitats. Anopsilana luciae differs further in the rounded
rather than angular anterior margin of the frontal process, which is visible in
dorsal view, and in the absence of marginal spines on the telson. Anopsilana
pustulosa differs in the presence of tubercles on the pleonites and posterior per-
eonites, the 2 carinae on the telson, and the more pointed telson. Anopsilana
willeyi also has a tuberculate body and a pointed telson with concave lateral
margins.

Anopsilana poissoni, like A. crenata, is a blind, unpigmented freshwater trog-
lobite, but has many points of difference, including a longer and narrower frontal
lamina, a pereopod | palm with 2 spines and no rounded scales, a much longer
appendix masculina with an abrupt bend, uropods much longer than the telson,
exopod longer than the endopod and much narrower than in A. crenata.

Habitat

Based on the geological map presented in Brunt et al. (1973), West Bay Cave
is developed in the limestone of the Ironshore Formation. This limestone, com-
posed of poorly consolidated reef limestone, calcarenites, and lagoonal muds and
sands, lies unconformably on the much older and more massive Bluff Limestone
Formation (Brunt et al. 1973). The Ironshore Formation, which occurs at ele-
vations of less than 5 meters above sea level, probably represents a Late Pleis-
tocene reef terrace. Other reef terraces from similar elevations in South Florida,
the Bahamas, and Jamaica have been dated at approximately 130,000 years BP

VOLUME 95, NUMBER 3 a1)

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Fig. 4. Known distribution of species of Anopsilana, except A. crenata: Solid stars, A. willeyi;
Open stars, A. pustulosa; Solid circle, A. luciae; Arrow, A. poissoni.

using Th??°/U2*4 ratios in unaltered fossil corals (Broecker and Thurber 1965, Cant
1972, Neumann and Moore 1975, Osmond et al. 1965). Presumably, the Ironshore
Formation is of similar age (Brunt et al. 1973, Morgan 1977).

The cave is located on the north flank of a low limestone ridge that rises
gradually from the present western shoreline. Access to the cave is through a
vertical sinkhole, approximately 2.5 meters deep. The cave itself consists of a
single chamber, about 8 meters in length, and 0.5—2.0 meters in height. The entire
chamber receives light from the entrance. The cave pool, which begins just inside
the entrance, covers the entire cave floor with up to 20 cm of fresh water. The
water is crystal clear and fresh to the taste. Presumably, as on other small, low-
lying Caribbean islands, the cave pool represents a locally-derived freshwater
lens atop a large volume of salt water. The floor of the cave, as well as the
adjacent sinkhole, consists of broken chunks of the white limestone; fine organic
silt is present in small areas adjacent to the entrance and the sinkhole. The silt
apparently is derived from surface soils and detritus funnelled through the sink-
hole into the cave. The sinkhole is shaded by gumbo-limbo (Bursera simaruba)
and West Indian almond (Terminalia catappa).

Cirolanid isopods were extremely abundant in the cave pool, as evidenced by
the large numbers of individuals represented in the sample. Swimming isopods
filled the water column, and were rarely observed at rest on the bottom or on the
vertical surfaces of the broken limestone. No other aquatic organisms were ob-
served in the cave pool. No bats, or signs of bats, were seen in the cave. Surface

528 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

animals such as amblypygids, scorpions, gastropods, and small geckoes (Sphae-
rodactylus argivus) were observed on the vertical walls and bottom of the sink
and on the slope adjacent to the water. Several adult treefrogs (Osteopilus sep-
tentrionalis) were found in small solution pits in the wall of the sink.

Zoogeography

Except for A. crenata the species of Anopsilana are distributed in the Indo-
Pacific (Fig. 4). Anopsilana willeyi is known from Lake Negombo, Ceylon (Steb-
bing 1904, Barnard 1935), Chilka Lake, India (Barnard 1935), and the coast of
Kerala state, India (Pillai 1954, 1961, 1967). Anopsilana pustulosa has been re-
ported from Cooktown, Queensland, Australia (Hale 1925), Tulear, Madagascar
(Roman 1970), and Aldabra atoll (Jones 1976). Anopsilana luciae is known only
from St. Lucia Bay, Zululand, South Africa (Barnard 1940). The type- and only
locality for A. poissoni, a cave in Madagascar, has already been mentioned.

Assigning these Indo-Pacific species to a common genus does not create prob-
lems in zoogeography, but the discovery of a congeneric freshwater troglobitic
species on a Caribbean island cannot be easily explained. Assigning the Cayman
Island species to Anopsilana implies that it shares common ancestry with the
Indo-Pacific species. The discovery of additional Atlantic species of Anopsilana
would be helpful, but cannot be anticipated.

A possible alternative explanation to common ancestry is convergence. The
only character separating Anopsilana from Cirolana is the absence of setae from
the endopods of pleopods 3-5. Bruce (1981) has pointed out that the species of
Anopsilana are from either brackish or freshwater habitats, and that reduction
of marginal setae is associated with movement into fresh water. Almost all fresh-
water Cirolanidae are troglobitic, and all 14 genera of troglobitic cirolanids lack
marginal setae on pleopods 3—5. Very few marine cirolanids have pleopods with
this attribute.

Thus it seems possible that at least some of the species of Anopsilana, and
especially A. crenata, have independently lost the setae of pleopods 3-5. If this
happened, the species involved do not have common ancestry and are not con-
generic. The convergence explanation is appealing for A. crenata, which can then
be derived from a local marine species of Cirolana, and its relation to the Indo-
Pacific species of Anopsilana need not be dealt with.

If it can be demonstrated convincingly that loss of the setae of pleopods 3-5
has evolved independently in the species of Anopsilana in response to decreased
salinity of their habitats, this genus will have to be abandoned and its species
returned to Cirolana. For the time being it is convenient to retain Anopsilana.

Acknowledgments
We thank C. W. Hart and Anne C. Cohen of the Smithsonian Institution for
reviewing the manuscript.

Literature Cited

Barnard, K. H. 1935. Report on the Amphipoda, Isopoda and Tanaidacea in the collections of the
Indian Museum.—Records of the Indian Museum 37:279-319.
1940. Contributions to the crustacean fauna of South Africa. 12. Further additions to the

VOLUME 95, NUMBER 3 529

Tanaidacea, Isopoda, and Amphipoda, together with keys for the identification of hitherto
recorded marine and freshwater species.—Annals of the South African Museum 32:381—515.

Broecker, W. S., and D. L. Thurber. 1965. Uranium-series dating of corals and oolites from Ba-
haman and Florida Keys limestones.—Science 149:58—60.

Bruce, N. L. 1981. Cirolanidae (Crustacea: Isopoda) of Australia: Diagnoses of Cirolana Leach,
Metacirolana Nierstrasz, Necirolana Hale, Anopsilana Paulian & Debouteville, and three new
genera—Natolana, Politolana and Cartetolana.—Australian Journal of Marine and Freshwater
Research 32:945-966.

Brunt, M. A., M. E. C. Gigliolo, J. D. Mather, D. J. W. Piper, and H. G. Richards. 1973. The
Pleistocene rocks of the Cayman Islands.—Geological Magazine 110(3):209-304.

Cant, R. V. 1972. Jamaica’s Pleistocene reef terraces.—Journal of the Geological Society of Jamaica
12:13-17.

Hale, H. M. 1925. Review of Australian isopods of the cymothoid group. Part I.—Transactions of
the Royal Society of South Australia 49: 128-185.

Jones, D. A. 1976. The systematics and ecology of some isopods of the genus Cirolana (Cirolanidae)
from the Indian Ocean region.—Journal of Zoology, London 178:209-222.

Monod, T. 1976. Remarques sur quelques Cirolanidés (Crustacés, Isopodes).—Bulletin du Muséum
National d’ Histoire Naturelle (3) 358, Zoologie 251:133-161.

Morgan, G. S. 1977. Late Pleistocene fossil vertebrates from the Cayman Islands.—Master’s Thesis,
University of Florida, Gainesville. 283 pp.

Neumann, C. A.,and W.S. Moore. 1975. Sea level events and Pleistocene coral ages in the northern
Bahamas.—Quaternary Research 5:215—224.

Osmond, J. K., J. R. Carpenter, and H. L. Windom. 1965. Th??°/U?*4 age of the Pleistocene oolites
and corals of Florida.—Journal of Geophysical Research 70(8): 1843-1847.

Paulian, R., and C. Delamare Deboutteville. 1956. Un Cirolanide cavernicole a Madagascar (Iso-
pode).—Memoires de |’Institut Scientifique de Madagascar série A, 11:85-88.

Pillai, N. K. 1954. A preliminary note on the Tanaidacea and Isopoda of Travancore.—Bulletin of
the Central Research Institute University of Travancore, Trivandrum 3(1):1—21.

———. 1961. Monograph wood-boring Crustacea of India——Government of India Press, Simla, x
+ 61 pp.

—. 1967. Littoral and parasitic isopods from Kerala: families Eurydicidae, Corallanidae and
Aegidae—2.—Journal of the Bombay Natural History Society 64(2):267—283.

Roman, M.-L. 1970. Ecologie et répartition de certains groupes d’Isopodes dans les divers biotopes
de la région de Tulear (sud-ouest de Madagascar).—Recueil des Travaux de la Station Marine
d’Endoume, Faculté des Sciences de Marseille, Fascicule Hors Série, Supplement no. 10:163-
208.

Stebbing, T. R. R. 1904. Marine crustaceans. XII. Isopoda, with description of a new genus.—
Fauna and. Geography of the Maldive and Laccadive Archipelagoes (J. Stanley Gardiner, ed.)
2(3):699-721, pls. 49-53.

Vandel, A. 1964. Biospéologie: la biologie des animales cavernicoles.—Gauthier- Villars, Paris, xviii
+ 619 pp., 11 pls.

(TEB) Department of Invertebrate Zoology (Crustacea), NHB 163, Smithson-
ian Institution, Washington, D.C. 20560; (RF) The Florida State Museum, Mu-
seum Road, University of Florida, Gainesville, Florida 32611.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 530-536

UNCISPIONIDAE, A NEW POLYCHAETE
FAMILY (ANNELIDA)

Karen D. Green

Abstract.—A new family of polychaetous annelids is described. This family
belongs in the order Spionida, and is most closely related to the Spionidae. Mem-
bers of this family have been reported from deep waters off Oregon, northeast
South America, and southern California (new record). Two genera are presently
known for the new family: Uncopherusa (reassigned) and Uncispio (new genus).
Uncispio hartmanae, a new genus and species, is described.

The Bureau of Land Management (BLM) sponsored a baseline study of the
Southern California Bight (contract no. AA550-CT6-40) during 1975-1978 (Fau-
chald and Jones 1978). From this study, several specimens of polychaetous an-
nelids that were collected in deep water off Santa Cruz Island were determined
by the author to be undescribed. This material, including some well preserved
and complete specimens, made it possible to establish the affinity of these worms
to spioniform families. The taxonomic placement of the proposed family within
the order Spionida is based on an analysis of characters used to define the spio-
niform families (Day 1967, Fauchald 1977, Hartman 1969).

Examination of the literature revealed two reports (Hartman 1965, Fauchald
and Hancock 1981) of polychaetes that belong within the proposed family. This
material is reassigned and compared to the new genus and species described here.

Material examined and type-specimens described in this paper are in the col-
lections of the Allan Hancock Foundation (AHF), University of Southern Cali-
fornia, Los Angeles, California.

Uncispionidae, new family

Diagnosis.—Small, slender polychaetes with palps inserted dorsally at junction
between pro- and peristomium (postectal prostomial margins). Occipital antenna
present. Parapodia biramous with reduced and simple lobes. Branchiae may be
present on a few antero-median segments, fused to notopodial lobes. Setae simple
and include capillaries (smooth or haired) and hooded bidentate hooks. Capillaries
long on first setiger, reminiscent of a cephalic cage; hooks become enlarged and
modified on a few posterior segments. Anus terminal, surrounded by four digitate
lobes.

Etymology.—The family name derives from the enlarged modified hooks (Latin
‘‘uncus’’ meaning hook) and affinity with the family Spionidae.

Remarks.—Uncispionids belong to the order Spionida Grube, 1850 and sub-
order Spioniformia because of the presence of a pair of grooved palps at the
postectal margins of the prostomium. A differential diagnosis (based on a suite
of characters) of each family within this suborder is presented in Table 1. Unci-
spionids share characters with poecilochaetids and trochochaetids (long capillar-
ies may be present on first setiger, haired setae may be present), heterospionids

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VOLUME 95, NUMBER 3

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VOLUME 95, NUMBER 3 533

ape A TNS OM

Fig. |. Uncispio hartmanae, holotype: a, Dorsal view, prostomium and first 2 setigers; b, Para-
type, dorsal view of head showing palp; c, Ventral view of anterior end; d, Dorsal view, pygidium
and last 3 setigers.

(reduced parapodial lobes, branchiae on thoracic setigers), but are most closely
related to spionids (hooded bidentate hooks, occipital antenna, branchiae). When
considered together, the presence of reduced parapodial lobes, haired setae (when
present), long capillaries on the first setiger, and modified hooded hooks precludes
placement of these worms into any previously described family.

534 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Key characters of type-material of Uncispio hartmanae.

Character Holotype Paratype Paratype
Length xX width 42x0.25mm 4.8 x 0.25mm _ 3.2 x 0.2 mm
Total number of setigers 39 37 31
Presence of branchiae; setigers 6-11 6-8 5-8
Presence of hirsute notosetae; setigers 15-31 9-31 11-23
Absence of notosetae; setigers 38—39 36-37 30-31
Presence of hooded neurohooks; setigers 9-39 9-37 8-31
Presence of modified neurohooks; setigers 38-39 36-37 30-31
Pygidial lobes 4 + cirrus 4 4
Palps absent absent present

In addition to the Santa Cruz Island specimens, members of this family have
been reported from deep waters (2860 m) off Oregon (as Uncopherusa bifida by
Fauchald and Hancock 1981) and northeast South America at a depth of 1500 m
(as ? Spionida by Hartman 1965).

Uncispio, new genus

Type-species.—Uncispio hartmanae, new species.

Diagnosis.—Body cylindrical with 3 distinct regions. Anterior setigers with
capillaries in noto- and neuropodia; first setiger with long bilimbate setae, other
setigers with short limbate setae with one edge finely haired. Median setigers
have in addition to short haired capillaries, long hirsute capillaries in notopodia
and bidentate hooded hooks in neuropodia. Posterior setigers without long hirsute
capillaries in notopodia; last 2 setigers with only neurosetae, hooded hooks en-
larged and modified.

Remarks.—Uncispio is related to Uncopherusa Fauchald and Hancock, 1981
and ? Spionida Hartman, 1965. Uncispio is differentiated from Uncopherusa by
the presence of haired notosetae (Uncopherusa lacks haired setae), and fewer
posterior setigers with modified hooded hooks (Uncopherusa has 5 setigers with
modified hooks). Uncopherusa was described as having the body partly covered
with papillae and encrusted with sand; it is suggested that this refers instead to
a description of the worm’s tube. Hartman’s specimen is recognized as belonging
to this family because of the presence of a similar pygidium and posterior modified
hooks. However, because the specimen is a fragment further taxonomic place-
ment is impossible.

Uncispio hartmanae, new species
Figs. 1, 2; Table 2

Material.—Holotype, AHF Poly 1365; paratypes, AHF Poly 1366; offshore
Santa Cruz Island, station 80901, 222 m, light brown hard clay with pebbles,
winter sample.

Description.—Holotype complete with 39 setigers, 4.2 mm in length, 0.25 mm
in width. Body lacking pigmentation in alcohol. Body cylindrical, segments more
elongate and less distinct in median than anterior and posterior regions of body.
Paratypes compared with holotype in Table 2. i

Prostomium anteriorly bluntly rounded with posterior occipital antenna; eyes

VOLUME 95, NUMBER 3 S35

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worm showing 3 body regions; b, Lateral view, fourth setiger; c, Lateral view, sixth setiger with
branchia; d, Lateral view, ca. setiger 31; e-f, Short limbate and hirsute capillaries, twelfth setiger
(ca. 500); g-1, Hooded hooks, setigers 25, 36 and 37, respectively (ca. 500).

absent (Fig. la). Peristomium forming lateral wings on either side of prostomium.
Palps missing on holotype, one palp present on paratype. Palps grooved and
short, inserted dorsally at junction between pro- and peristomium lateral to the
occipital antenna (Fig. 1b). Proboscis an axial sac with what appears to be 2 distal

536 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

digitate lobes. However, because proboscis is not completely everted, and the
worm is small, it is difficult to distinguish if these lobes are actually part of the
proboscis or subdistal antennae (Figs. la, b, c).

Anus terminal, surrounded by 4 primary digitate lobes, pair ventral and dorsal;
holotype with one additional smaller cirrus (Fig. 1d).

Parapodia biramous with reduced rounded lobes. All setae simple, include cap-
illaries and hooded bidentate hooks. Setae distinct in anterior, median and pos-
terior regions of body (Fig. 2a). First setiger with very long bilimbate capillaries
in noto- and neuropodia; setae splayed, surround peristomium, reminiscent of
cephalic cage. Setigers 2-8 with short limbate (with one edge finely haired) cap-
illaries, approximately 6 per fascicle, both in noto- and neuropodium (Figs. 2b,
c, e). Branchiae fused to notopodial lobes (Fig. 2c), present on setigers 6-11.

Noto- and neurosetae of median segments include in addition to short haired
capillaries, long whip-like hirsute capillaries in notopodium (1 hirsute and 1-2
short capillaries per fascicle), and hooded bidentate hooks in neuropodium (2
capillaries and 1-3 hooks per fascicle) (Figs. 2d, f, g). Hirsute capillaries are
present in setigers 15-31. However, presence of hirsute capillaries coincident
with presence of hooded hooks in paratype may indicate that these setae were
missing from more anterior setigers on the holotype. Hooded hooks are present
in the neuropodium of setiger 9 and continue to end of worm.

Notosetae of posterior setigers (setigers 32—39) with only short haired capillar-
ies, similar to anterior setigers (2 per fascicle). Last 2 setigers (38-39) lack no-
tosetae; neurohooks enlarged and modified (penultimate hook largest), remnant
of hood present (Figs. 2h, 1).

Etymology.—The species is named in honor of the late Dr. Olga Hartman who
first discovered this family and recognized its affinity to spionids.

Distribution.—Offshore Santa Cruz Island, California, 222 m.

Acknowledgments

I would like to thank Drs. Kristian Fauchald, James A. Blake, and Jerry D.
Kudenov who shared some of their thoughts regarding the taxonomic placement
of these unusual worms. My thanks also extend to Dr. John H. Dorsey and Dr.
Kristian Fauchald for reviewing and commenting on the manuscript.

Literature Cited

Day, J. H. 1967. A monograph of the polychaetes of southern Africa, part 2: Sedentaria.—The
British Museum (Natural History) Publication 656: 1-878.

Fauchald, K., and D. R. Hancock. 1981. Deep-water polychaetes from a transect from off central

Oregon.—Monographs of the Allan Hancock Foundation 11:1-73.

, and G. F. Jones. 1978. Variation in community structure of shelf, slope and basin macro-
Panna communities of the southern California Bight.—Report from SAI (contract no. AA550-
CT6-40) to BLM, Dept. of Interior.

———. 1977. The polychaete worms: definitions and keys to the orders, families and genera.—
Natural History Museum of Los Angeles County, Science Series 28:1—-188.

Hartman, O. 1965. Deep-water benthic polychaetous annelids off New England to Bermuda and
other North Atlantic areas.—Allan Hancock Pacific Expedition 19:187—456.

——. 1969. Atlas of sedentariate polychaetous annelids from California.—Allan Hancock Foun-
dation, University of Southern California, Los Angeles. 812 pp.

3513 Ponderosa Drive, Oceanside, California 92054.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 537-544

FOUR NEW SPECIES OF STOMATOPOD CRUSTACEANS
FROM THE PHILIPPINES

Renato G. Garcia and Raymond B. Manning

Abstract.—Haptosquilla philippinensis, Clorida nazasaensis, Levisquilla ar-
mata, and Oratosquilla microps, all from localities in the Philippine Islands, are
named.

The species reported here were discovered by one of us (R.G.G.) during a
preliminary survey of the stomatopod fauna of the Philippines. Holotypes of the
four new species are deposited in the National Museum of the Philippines, Manila
(NMCR); some paratypes also have been deposited in the Crustacea collection,
National Museum of Natural History, Smithsonian Institution, Washington, D.C.
(USNM). All measurements are in millimeters.

We thank Lilly King Manning for preparing the illustrations.

Family Protosquillidae Manning, 1980
Haptosquilla philippinensis, new species
Je I

Material.—Talaan, Sariaya, Quezon, Tayabas Bay, 1345’N, 121°45’E; J. Sy
and R. Rivera, leg.; March 1978: 1 female (holotype, NMCR 2866).

Description.—Cornea subglobular, set obliquely on stalk. Ocular scales well
developed, produced into triangular lobes laterally, falling short of lateral rostral
spines.

Rostral plate trispinous, median spine more slender and longer than laterals,
latter slender, sharp, extending to bases of eyestalks.

Anterior margins of lateral plates of carapace slightly concave, anterolateral
angles rounded, not strongly produced.

Mandibular palp 2-segmented.

Inflated part of outer margin of dactylus notched. Propodus with single movable
spine proximally on inner margin.

Anterior 4 abdominal somites smooth, unarmed, not carinate dorsally. Fifth
somite smooth medially, with 2 low longitudinal carinae laterally above lateral
margin, separated by a deep groove. Sixth somite with 6 carinae, each spined
posteriorly, submedians and intermediates rather broad, inflated, ornamented lat-
erally with scattered setae.

Telson slightly broader than long, with 4 pairs of marginal teeth, submedians
with movable apices. Dorsal surface of telson with 3 bosses, median smallest,
triangular, laterals longer, oblong, extending about to midlength of telson but
beyond median cleft. All 3 bosses ornamented with setae. Inner margins of sub-
median marginal teeth with 8 denticles, intermediate and lateral teeth each with
fixed mesial denticle, marginal teeth lacking denticles.

Uropods stout, proximal segment of exopod with 9 short movable spines, dis-
talmost extending beyond midlength of distal segment. Inner spine of basal pro-
longation of uropod well developed but shorter than outer.

538 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Haptosquilla philippinensis, female holotype: a, Anterior part of body; b, Uropod, ventral
view; c, Sixth abdominal somite, telson, and uropod.

Color.—Body ornamented with dark chromatophores in 2 broad bands across
carapace and scattered over thoracic and abdominal somites. Sixth thoracic so-
mite with broad median dark patch, extending posteriorly onto seventh somite,
as well as a pronounced dark spot on each lateral process. Second abdominal
somite with dark rectangular patch, fifth with submedian pair of rounded dark
spots.

Measurements .—Total length 20; carapace length 4.0; eye length 2.1; cornea
width 0.9; rostral plate length 1.6, width 1.9; fifth abdominal somite width 3.8;
telson length 3.4, width 3.7.

Remarks.—This small species resembles H. proxima Kemp (1915: 183, pl. 1,
figs. 9, 10) in most respects, but differs in several features: the mandibular palp
is present, the bosses of the sixth abdominal somite are armed posteriorly, and
the dorsal bosses of the telson are larger and are shaped differently. In H. proxima
all three bosses are subequal in size and almost circular in shape, whereas in H.
philippinensis the median is triangular and the laterals are larger and oval in shape.

Haptosquilla philippinensis also resembles H. setifera Manning (1969:162, fig.
8) from Bougainville, Solomon Islands, but in the latter species the median boss
of the telson is comparatively larger, resembling the laterals in size and shape.

Etymology.—The specific epithet reflects the location of the type-locality in
the Philippine Islands.

Family Squillidae Latreille, 1803
Clorida nazasaensis, new species
Fig. 2

Material.—Nazasa Bay, Zambales, 14°49’N, 120°06’E; 9-20 fms [3-37 ml];
dredge; F. G. Dayrit and J. E. Norton, Leg.; 30 January—8 February 1960: 1 male
(holotype, NMCR 741).

Description.—Size small, total length of adult male 43 mm.

Eye large, flask-shaped, extending about to middle of first segment of anten-
nular peduncle. Cornea small, bilobed, set obliquely on stalk, cornea width about
¥ eye length, 2 stalk width. Ocular scales fused into bilobed plate.

Antennular peduncle as long as carapace.

Rostral plate short, triangular, broader than long, median carina absent.

VOLUME 95, NUMBER 3 539

Fig. 2. Clorida nazasaensis, male holotype: a, Anterior part of body; b, Claw; c, Lateral processes
of exposed thoracic somites; d, Sixth abdominal somite, telson, and uropod; e, Uropod, ventral view.

Carapace strongly narrowed anteriorly, anterior width less than half median
length. Surface smooth, lacking carinae except for reflected marginals on post-
erior part of lateral plates. Anterolateral spines strong but falling short of base of
rostral plate.

Mandibular palp absent; 4 epipods present.

Raptorial claw large, stout. Dactylus slender, with 4 teeth on inner margin,
proximal very small, appressed to base of next tooth. Propodus strongly inflated,
opposable margins pectinate on proximal *%4 of outer margin, with 3 movable
spines proximally on inner margin. Dorsal ridge of carpus undivided, ending in
blunt lobe ornamented mesially with patch of setae.

Exposed thoracic somites lacking submedian carinae, posterior 3 somites with
intermediate carinae. Lateral process of fifth thoracic somite short, appearing
acute in dorsal view, in lateral view a rounded lobe, compressed antero-post-
eriorly. No ventral spine present on fifth somite. Lateral processes of sixth and
seventh somites broadly rounded laterally.

Abdomen broad, strongly depressed, submedian carinae absent except on sixth
somite. Abdominal carinae spined as follows: submedian 6, intermediate 5-6,
lateral 6, marginal 5.

Telson broader than long, inflated, with 3 pairs of marginal teeth, submedians

540 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

with movable apices. Lateral margins of intermediate and lateral teeth tuberculate
or crenulate. Short, inconspicuous prelateral lobes present. Dorsal surface of
telson with numerous broad tubercles, as illustrated. Denticles spiniform, 3-4, 7,
1. Ventral surface lacking distinct postanal Keel.

Uropods stout, proximal segment of exopod shorter than distal, with 5 movable
spines on outer margin, proximal 3 small, spiniform, situated dorsally, distal 2
much larger, situated marginally, distalmost spatulate, extending about to mid-
length of distal segment. Inner spine of basal prolongation of uropod the longer,
with low, rounded lobe on outer margin and 6 strong spines on inner margin.

Color.—Carapace outlined with thin line of dark pigment, similar thin line
present posteriorly on posterior 3 thoracic and anterior 5 abdominal somites.
Uropod with a dark spot on inner distal margin of proximal segment of exopod,
distal segment dark on inner half.

Measurements.—Male holotype, total length 43 mm. Other measurements: car-
apace length 9.0; anterior width carapace 4.1; cornea width 0.7; eyestalk length
2.2, width 1.4; antennular peduncle length 9.0; rostral plate length 1.0, width 1.7;
telson length 7.5, width 10.2.

Remarks.—This new species most closely resembles C. seversi Moosa (1973:
22, fig. 4), a species described from Indonesia. It differs from the latter species
in several important respects: there are only 3 rather than 4 well formed teeth on
the dactylus of the claw, the proximal being very reduced; there are fewer, larger
dorsal teeth on the telson and no postanal keel ventrally; and there are only 5
movable spines on the uropod, the proximal 3 slender, the distal 2 much enlarged,
with the distalmost spatulate.

Etymology.—The specific epithet is derived from the type-locality.

Levisquilla armata, new species
Fig. 3

Material.—Cape Calabite, Occidental Mindoro, 13°23’N, 120°19’E; F. G. Dayrit
and J. E. Norton, leg.; March 1960: 16 males, 14 females (NMCR 802; 2 males,
2 females from this lot USNM 190708).—Off Laoy Island, Bohol, 09°36’N, 124°01’E;
45 fms (82 m); dredge; Pele-Sulu Sea Expedition; 8 February 1964: 3 males, 2
females (NMCR 1370).—South Panglao, Panglao Island, Bohol, 09°35’N, 123°48’E;
45-70 fms (82-128 m); dredge; Pele-Sulu Sea Expedition: 2 males (NMCR 1384;
larger male is holotype, remainder of specimens are paratypes).

Description.—Size small, total length of adults less than 50 mm. Dorsal surface
of body appearing smooth, polished.

Eye small, elongate, cornea bilobed, set obliquely on stalk. Eyes extending to
or almost to end of first segment of antennular peduncle. Corneal indices 338—
500, lower in smaller specimens.

Antennular peduncle slightly shorter than carapace. Dorsal processes of an-
tennular somite produced into acute projections directed anterolaterally.

Rostral plate triangular, length and width subequal or slightly longer than broad,
without carinae, apex sharp.

Carapace smooth, anterior width about half median length. Anterolateral spines
strong but falling short of base of rostral plate. Median carina completely absent,
short intermediates and reflected marginals present posteriorly on lateral plates.

VOLUME 95, NUMBER 3 54]

Fig. 3. Levisquilla armata, female paratype, TL 38 mm: a, Anterior part of body; b, Eye; c,
Claw; d, Lateral processes of fifth, sixth, and seventh thoracic somites; e, Fifth and sixth abdominal
somites, telson, and uropod; f, Uropod, ventral view.

Mandibular palp absent; 4 epipods present.

Dactylus of claw with 6 teeth, outer margin evenly curved or faintly flattened,
with strong basal notch. Dorsal ridge of carpus undivided. Inferodistal angle of
merus unarmed.

Exposed thoracic somites lacking submedian carinae, intermediates present on
posterior 3 somites. Lateral process of fifth thoracic somite a slender spine di-
rected anterolaterally, flanked ventrally by a slender spine. Lateral processes of
sixth and seventh somites undivided, rounded, that of seventh somite the larger.

Abdomen lacking any trace of submedian carinae on anterior 5 somites, re-
mainder of carinae spined as follows: submedian 6, intermediate 5—6, lateral 5—
6, marginal (3-4)-—5.

Telson broader than long, with well marked median carina, short carinae on
marginal teeth, and prelateral lobe. Dorsal surface with several curved rows of
short carinae or rows of tubercles on each side, as illustrated. Submedian teeth
with movable apices. Denticles sharp and slender, 3-5, 9-13, 1 (usually 4, 10-11,
1). Ventral surface with short but distinct median carina.

542 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Uropod rather stout, proximal segment of exopod (measured dorsally) shorter
than distal, with row of 7—9 graded spines on outer margin, proximals slender,
sharp, distal 2 enlarged, spatulate. Basal prolongation with 2 rounded iobes be-
tween apical spines and with row of 5—13 (usually 9) slender spines along inner
margin.

Color.—Completely faded.

Measurements.—Total lengths of males 21 to 44.5 mm, of females 22 to S0
mm. Other measurements of male holotype, 43.5 mm: carapace length 9.2; an-
terior width of carapace 4.5; antennular peduncle length 8.8; cornea width 2.0;
rostral plate length 1.5, width 1.4; telson length 7.4, width 8.2.

Remarks.—This new species can be distinguished immediately from the only
other species assigned to Levisquilla (see Manning 1977:422), the type-species L.
inermis (Manning, 1965:255, fig. 2) by the dorsal carinae on the telson; in L.
inermis the dorsal surface of the telson is smooth. Further, in L. inermis the
rostrum is more rounded anteriorly, the dorsal processes of the antennular somite
are more slender and sharper, and the proximal of the 2 lobes between the spines
of the basal prolongation of the uropod is much less pronounced.

Etymology.—The specific epithet is from the Latin, armatus, armed.

Oratosquilla microps, new species
Fig. 4

Material.—Cabulan Island, Bohol, 10°10’N, 124°02’E; 20-30 fms (37-55 m);
dredge; Pele-Sulu Sea Expedition; 5 February 1964: 1 male (holotype, NMCR
2868).

Description.—Size small, total length of adult male holotype 50 mm. Body
appearing smooth, lightly pitted under magnification.

Eye very small, cornea bilobed, set obliquely on stalk. Eyes not extending to
end of first segment of antennular peduncle. Corneal index 492.

Antennular peduncle subequal to carapace in length. Dorsal processes of an-
tennular somite produced into acute spines directed anterolaterally.

Rostral plate tongue-shaped, slightly broader than long, apex rounded. Median
carina absent. Anterior width of carapace more than half median length. Antero-
lateral spines strong but falling short of base of rostral plate. Median carina
lacking anterior bifurcation. Intermediate carinae falling short of anterior margins
of lateral plates.

Mandibular palp present; 4 epipods present.

Dactylus of claw with 6 teeth, outer margin sinuate. Dorsal ridge of carpus
undivided. Inferodistal angle of outer face of merus with acute spine.

Exposed thoracic somites with unarmed submedian and intermediate carinae.
Lateral process of fifth somite bilobed, anterior lobe a slender, sharp, anteriorly
directed spine, posterior lobe smaller, triangular. Lateral process of sixth somite
bilobed, anterior lobe very slender, rectangular, distally truncated, blunt, pos-
terior lobe triangular, apex acute but rounded. Lateral process of seventh somite
distinctly bilobed, anterior lobe much smaller than posterior, apex of latter almost
forming a right angle.

Submedian carinae slightly divergent on abdominal somites, especially poste-
riorly. Abdominal carinae spined as follows: submedian 5—6, intermediate 4-6,
lateral 2-6, marginal 1-5.

VOLUME 95, NUMBER 3 543

Fig. 4. Oratosquilla microps, male holotype: a, Anterior part of body; b, Eye; c, Carpus of claw;
d, Lateral processes of exposed thoracic somites; e, Fifth and sixth abdominal somites, telson, and
uropod; f, Posterior margin of telson, ventral view; g, Uropod, ventral view.

Telson flattened, slightly longer than broad, dorsal surface ornamented with
numerous carinae and tubercles in curved rows, as illustrated. Prelateral lobe
longer than margin of lateral tooth. Denticles rounded, 3, 9, 1, many with minute
apical spinule. Ventral surface with long postanal keel.

Uropod slender, long, proximal segment of exopod longer than aictall with 8
movable spines on outer margin, distalmost not extending to midlength of distal
segment. Lobe on outer margin of inner spine of basal prolongation rounded,
about as wide as adjacent spine, margin concave.

Color.—Completely faded.

Measurements.—Total length of male holotype 50 mm; other measurements:
carapace length 11.8; anterior width carapace 6.4; length antennular peduncle
11.5; cornea width 2.4; rostral plate length 1.8, width 2.1; telson length 10.2,
width 9.4.

Remarks.—This new species is a member of the perpensa complex of the per-
pensa group of species within the genus Oratosquilla (see Manning 1978:4, for a
discussion of species groups in Oratosquilla), those species in which the infe-
rodistal angle of the outer face or the merus of the claw is armed, the dactylus

544 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

of the claw has six teeth, the ridge on the carpus of the claw is undivided, and
the rostral plate is short. The dorsal carinae of the telson will immediately dis-
tinguish Oratosquilla microps from the three species now comprising that com-
plex, O. anomala (Tweedie), O. perpensa (Kemp), and O. stephensoni Manning.
Further, both O. anomala and O. perpensa have distinct anterior branches of
the median carina of the carapace, which are completely absent in O. microps,
and in O. stephensoni, a much larger species (total length to 150 mm), the lobe
between the spines of the basal prolongation of the uropod is convex or straight,
rather than concave.

Etymology.—The specific epithet is from the Latin and alludes to the relatively
small eyes of this species.

Literature Cited

Kemp, S. 1915. Ona collection of stomatopod Crustacea from the Philippine Islands.—The Phil-
ippine Journal of Science 10(3D):169—187, plate 1.

Manning, Raymond B. 1965. Stomatopoda from the collection of His Majesty The Emperor of

Japan.—Crustaceana 9(3):249-262, figures 1, 2, plates 11, 12.

1969. Notes on the Gonodactylus section of the family Gonodactylidae (Crustacea, Sto-
matopoda), with descriptions of four new genera and a new species.—Proceedings of the
Biological Society of Washington 82:143—166, figures 1-8.

—. 1977. Preliminary accounts of five new genera of stomatopod crustaceans.—Proceedings
of the Biological Society of Washington 90(2):420-423.

—. 1978. Further observations on Oratosquilla, with accounts of two new genera and nine new
species (Crustacea: Stomatopoda: Squillidae)—Smithsonian Contributions to Zoology 272: 1-—
44, figures 1-25.

Moosa, M. Kasim. 1973. The stomatopod Crustacea collected by the Mariel King Memorial ex-
pedition in Maluku waters in 1970.—Penelitian laut di Indonesia [Marine Research in Indonesia]
13:1-30, figures 1-4.

(R.G.G.) Carcinology Section, Zoology Division, National Museum, P.O. Box
2659, Manila, Republic of the Philippines; (R.B.M.) Department of Invertebrate
Zoology, National Museum of Natural History, Smithsonian Institution, Wash-
ington, D.C. 20560, U.S.A. |

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 545-553

A NEW CRAYFISH OF THE GENUS PROCAMBARUS FROM
SOUTHWESTERN ARKANSAS

Horton H. Hobbs, Jr., and Henry W. Robison

Abstract.—Procambarus (Girardiella) parasimulans is described from the
Ouachita and Red river basins of southwestern Arkansas. Its relationships to P.
(G.) curdi Reimer and P. (G.) simulans (Faxon) are discussed.

Comparatively little attention has been accorded the crayfishes of Arkansas
occurring south of the Arkansas River basin. The review of Bouchard and Rob-
ison (1981) summarizes the currently known fauna, but details of the distribution
of few of the species have been recorded by anyone. For obvious reasons, the
primary and secondary burrowers belonging to the genus Fallicambarus and to
the subgenus Girardiella of the genus Procambarus are poorly represented in
collections, and first form males are rare among the few series that are available.
The species described herein first came to our attention in a single collection
made in Sevier County in 1973 (see ‘“‘Range and specimens examined’’); however,
not until subsequent field work by one of us (H.W.R.) resulted in obtaining ad-
ditional material were the available specimens considered adequate for preparing
a description. Moreover, in view of the unpublished study by Rollin D. Reimer,
we were and remain hesitant to describe crayfishes that almost certainly had been
recognized as undescribed by him. In view of no response to our attempts to
communicate with Dr. Reimer, however, we offer this description.

Procambarus (Girardiella) parasimulans, new species
Fig. 1

Diagnosis.—Body pigmented, eyes well developed. Rostrum of adults without
marginal spine and median carina. Carapace lacking cervical spine or tubercle.
Areola 5.1 to 9.0 (average 6.6 + 0.9822) times as long as broad and constituting
30.0 to 35.0 (average 32.5) percent of total length of carapace (37.8 to 42.3 (av-
erage 39.9) percent of postorbital carapace length). Suborbital angle weak, lacking
spine or tubercle in adult. Hepatic area weakly tuberculate; branchiostegal spine
reduced to weak tubercle. Antennal scale about twice as long as broad, widest
at, or slightly distal to, midlength. Ventral surface of chela strongly tuberculate,
tubercles present along proximal half of ventral surface of dactyl. Ischium of third
pereiopod of first form male with simple, strong hook overreaching basioischial
articulation; hook not opposed by tubercle on corresponding basis; coxa of fourth
pereiopod lacking caudomesial boss. First pleopods of first form male reaching
coxae of third pereiopods, symmetrical, bearing proximomesial spur at caudal
proximomesial angle and subtruncate cephalic shoulder at base of terminal ele-
ments, lacking subterminal setae; terminal elements (all at least partly cornified)
consisting of (1) long, slightly curved mesial process reaching beyond other ele-
ments distally; (2) small, weakly curved, cephalodistally directed cephalic process
at cephalic base of mesial process; (3) similarly disposed, strongly cornified,

546 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

clawlike central projection arising from between mesial and caudal processes;
and (4) very conspicuous caudal process rounded, spatulate, and disposed ceph-
alodistally. Female with annulus ventralis about 1.5 times as broad as long, sub-
symmetrical in outline, with curved cephalomedian trough, flanked by rows of
prominent tubercles (or at least by a scalloped wall), leading to sigmoid sinus,
latter ending near median line anterior to caudal margin of annulus; preannular
plate not recognizable; postannular sclerite about 0.7 as wide and little more than
0.5 as long as annulus, broadly rounded anteriorly and weakly elevated ventrally;
first pleopod present.

Holotypic male, form I: Cephalothorax (Fig. la, 0) subovate, weakly com-
pressed laterally; maximum width of carapace slightly greater than height at cau-
dodorsal margin of cervical groove (17.8 and 17.3 mm). Abdomen narrower than
thorax (14.1 and 17.8 mm). Areola 7.1 times as long as wide, with | to 3 punc-
tations across narrowest part. Cephalic section of carapace about 1.8 times as
long as areola, latter comprising 34 percent of total length of carapace (41.3
percent of postorbital carapace length). Surface of carapace punctate dorsally
except in polished gastric area anterior to articular level of cardiac stomach,
granulate laterally, and weakly tuberculate in hepatic region. Rostrum broad with
weakly convergent margins turning suddenly anteromesially over penultimate
podomere of antennular peduncle and ending in slightly upturned tip at base of
ultimate podomere of peduncle; margins not conspicuously thickened; upper sur-
face concave with submarginal rows of punctations, very few others except clus-
ters of several larger ones on lateral sides of median depression at rostral base.
Subrostral ridge weak, barely perceptible except along caudal margin of orbit.
Postorbital ridge rather weak, neither swollen caudally nor with tubercle or spine
cephalically. Suborbital angle and branchiostegal spine both weak, latter con-
sisting of hardly more than very small tubercle and only slightly larger than those
tubercles flanking ventral side of cephaloventral part of cervical groove. Cervical
Spine absent.

Abdomen only little longer than carapace (36.3 and 35.6 mm). Pleura of third
through fifth segments subtruncate ventrally and rounded caudoventrally. Ce-
phalic section of telson with 4 spines (that next to lateralmost movable) in each
caudolateral corner (Fig. 1k). Cephalic lobe of epistome (Fig. Im) subtriangular
with weakly undulating, somewhat thickened, anterolateral margins; median sur-
face arched ventrally; main body of epistome with subtriangular depression mark-
ing area usually occupied by median fovea; epistomal zygoma broadly arched.
Ventral surface of proximal podomere of antennular peduncle with heavy short
spine just proximal to midlength. Antennal peduncle with short distolateral spine
on basis; ischium without ventral spine or tubercle; flagellum broken but reaching
base of fifth abdominal segment. Antennal scale (Fig. In) about twice as long as
broad, widest at about midlength; greatest width of lamellar area almost 3 times
that of thickened lateral part.

Third maxilliped extending as far anteriorly as antennal peduncle, distinctly
overreaching antennular peduncle; mesial sector of ventral surface of ischium
with crowded long stiff setae, lateral sector with submarginal row and scattered
short, more delicate plumose setae; merus similarly clothed.

Right chela (Fig. Ir) subovate in cross-section, somewhat depressed; palm
approximately 1.2 times as broad as length of mesial margin; latter almost one-

VOLUME 95, NUMBER 3 547

Fig. 1. Procambarus (G.) parasimulans (a, b, d-f, h-o, and r from holotype; c, g from morpho-
type, and p, q from allotype): a, Lateral view of carapace; b, c, Mesial view of first pleopod; d,
Cephalic view of distal part of first pleopod; e, Caudal view of first pleopods; f, Caudal view of distal
part of first pleopod; g, h, Lateral view of first pleopod; i, Mesial view of distal part of first pleopod;
J, Lateral view of distal part of first pleopod; k, Dorsal view of telson and uropods; /, Ventral view
of basal podomeres of third, fourth, and fifth pereiopods; m, Epistome; n, Antennal scale; 0, Dorsal
view of carapace; p, Annulus ventralis and adjacent sternal features; g, r, Dorsal view of distal
podomeres of first pereiopod.

548 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

third total length of chela; except for narrow ventrolateral area and ventro- and
dorsomedian ridges on fingers, surface studded with squamous to subsquamous
tubercles. Mesial margin of palm with row of 7 (left with 8) tubercles flanked
dorsally and ventrally by 2 sublinear series and few additional tubercles. Both
fingers with low median longitudinal ridges dorsally and ventrally; ridges flanked
proximally by squamous tubercles and distally by punctations. Opposable margin
of fixed finger with row of 15 (left with 10) tubercles (fourth, third on left, from
base largest) of which some of more distal ones so small that not depicted in
illustration; row reaching base of distal third of finger; large, subtriangular tu-
bercle present on lower level at base of distal fourth of finger; opposable margin
also bearing row of minute denticles, interrupted by tubercles, extending from
seventh tubercle from base of finger to proximal end of corneous tip of finger.
Opposable margin of dactyl distinctly excised proximally, bearing row of 19 (left
with 16) tubercles (fourth from base largest) along proximal two-thirds, and mi-
nute denticles opposing those on fixed finger; mesial margin of dactyl with row
of 10 tubercles along proximal two-thirds of finger, tubercles decreasing in size
and becoming more squamous toward distal end of finger.

Carpus of cheliped longer than broad with sinuous furrow dorsally; furrow
flanked by tubercles mesially and distolaterally and punctations proximolaterally;
mesial surface of podomere with 5 (left with 7) tubercles, that near midlength
largest, ventromesial surface with cluster of tubercles and ventrodistal margin
with usual 2 tubercles, none lateral to that serving as condyle ventrolaterally.

Merus with dorsal tubercular band extending from base almost to distal end of
podomere, band originating in single row and broadening and becoming generally
more conspicuous distally; mesial and lateral surfaces comparatively smooth;
ventral surface with mesial row of 17 (left with 15) tubercles, lateral row of 8 (left
with 7), and distal oblique row of 3 joining mesial and lateral rows; 3 small
tubercles in row along lateral side of distal articular membrane of podomere and
another small one at base of distolateral condyle. Ischium with row of 6 (left with
5) tubercles ventromesially.

Hook on ischium of third pereiopod (Fig. 1/) simple, heavy, thumblike, and
overreaching basioischial articulation, not opposed by strong tubercle on basis
but rudiment of one present on basis of left pereiopod. Coxa of neither fourth
nor fifth pereiopods with boss, but ventral caudomesial angle of fifth slightly
produced.

Sternum between third, fourth, and fifth pereiopods comparatively shallow but
ventrolateral margins bearing fringe of plumose setae, latter not concealing first
pleopods.

First pleopods (Fig. 1b, d-f, h-j) as described in ‘“‘Diagnosis.’’ In addition,
proximomedian lobe moderately long.

Uropods (Fig. 1k) with both lobes of proximal podomere bearing short, acute
spines; mesial ramus with distomedian spine small and situated distinctly pre-
marginally.

Allotypic female: Differing from holotype, other than in secondary sexual fea-
tures, in following respects: rostrum with margins more strongly convergent and
acumen more distinctly delimited; branchiostegal spine more spiniform but hardly
more conspicuous; cephalic section of telson with only 2 spines in each caudo-
lateral corner, more mesial of which movable; third maxilliped overreaching an-

VOLUME 95, NUMBER 3 549

Table 1.—Measurements (mm) of Procambarus (G.) parasimulans.

Holotype Allotype Morphotype

Carapace:

Entire length 35.6 38.9 28.7

Postorbital length 2933 322 23.7

Width 17.8 19.2 42

Height ES 18.4 13.4
Areola:

Width Mey De 15

Length Weil 12.8 9.6
Rostrum:

Width 6.1 6.3 4.6

Length 8.1 8.2 6.1
Chela:

Length, palm mesial margin 11.3 7.8 6.1

Palm width 13.3 10.1 7.6

Length, lateral margin 32.9 24.4 18.4

Dactyl length 20.0 14.7 les
Abdomen:
~ Width 14.1 16.3 10.9

Length 36.3 40.8 2B), 1|

tennal peduncle by length of distal podomere, ischium and merus much less
strongly hirsute; mesial margin of palm of chelae (Fig. 1q) with row of 8 tubercles,
opposable margin of fixed finger of chela with row of 11 (13 on left) tubercles,
row of minute denticles originating at distal base of first tubercle in row and
becoming band for short distance beyond triangular tubercle on lower level; op-
posable margin of dactyl of chela with row of 14 (12 on left) tubercles and row
of minute denticles originating immediately distal to tubercle marking distal mar-
gin of excised area of finger; mesial surface of carpus of cheliped with row of 7
tubercles; ventral surface of merus with mesial row of 15 (17 on left) tubercles
and lateral one of 8, oblique row of 2 (right) or 3 (left), and only 2 in row flanking
articular membrane of right chela; ischium with row of 4 (right) or 5 (left) tuber-
cles.

Annulus ventralis (Fig. lp) as described in “*‘Diagnosis.”’

Morphotypic male, form II: Differing from holotype in following respects: bran-
chiostegal spine distinctly spiniform; cephalic section of telson with 2 spines in
each caudolateral corner, more mesial one movable; shallow median fovea pres-
ent in main body of epistome; ischium of antennal peduncle with small tubercle
on ventral surface; antennal flagellum reaching third abdominal tergum; antennal
scale broadest slightly anterior to midlength; third maxilliped reaching very little
beyond antennal peduncle; mesial margin of palm of chela with row of 7 tubercles;
opposable margin of fixed finger with row of 10 tubercles (third from base largest)
and that of dactyl with 13 (right) or 12 (left) tubercles, fourth from base largest;
mesial surface of carpus with 6 tubercles; ventral surface of merus with lateral
row of 9 (right) or 10 (left) tubercles, mesial row of 15 (right) or 19 (left), and
oblique row of 4 tubercles; ventromesial margin of ischium with row of 4 tuber-

550 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

cles; ischium of third pereiopod tuberculiform and not attaining basioischial ar-
ticulation; sternum between third, fourth, and fifth pereiopods shallow, and setae
on ventrolateral margins not so well developed.

First pleopod (Fig. Ic, g) with 4 elements represented but none corneous, and
juvenile suture evident. All terminals more robust but disposed as in first form
male.

Color notes.—(Based upon specimens from Hot Spring and Sevier counties,
Arkansas.) Carapace pale tan dorsally fading to cream ventrolaterally. Dark brown
stripe on ventral flank of postorbital ridge and another extending ventrally across
orbital and antennal areas, setting off marginal, subtriangular cream marking on
antennal and upper anteroventral branchiostegal regions. Hepatic area with paler
brown reticulations; pale to dark brown pattern covering most of paired mandib-
ular adductor regions, these patches joined by narrow dark band along antero-
median margin of cervical groove. Branchiostegites with large, dark brown areas
dorsolaterally and somewhat paler brown reticulate stripe more ventrally, latter
extending from cervical groove almost to posterior margin of carapace. Abdomen,
like carapace, tan dorsally (however, some specimens with tergum of first seg-
ment dark brown) fading ventrally to very pale tan and marked by 2 pairs of
scalloped dark stripes: more dorsal one darker and extending caudally from level
of upper patch on branchiostegite and terminating on sixth tergum; more ventral
stripe, corresponding in level to ventral stripe on branchiostegite, consisting of
ventrally convex arcs along bases of pleura; latter with cream to white spots.
Uropods tan with brown reticulations, and lateral ramus with dark brown lateral
border. Antennules mostly dark brown; antennal peduncle tan with dark splotch-
es, and lateral border of antennal scale also quite dark. Dorsal surface of cheliped
much darker than ventral, dark brown reticulations most conspicuous along dor-
sal edge and on distal border of merus, and only slightly less so on dorsum of
carpus, on dorsomesial part of palm, and on dorsal surface of dactyl; dark col-
oration becoming more dilute laterally on both palm and fixed finger of chela.
Remaining pereiopods, especially third and fourth, with brown reticulations dor-
sally from ischium almost to distal extremity of propodus, darkest on contiguous
parts of merus and carpus, from which fading proximally and distally.

Type-locality.—An unnamed tributary to Prairie Bayou (Ouachita River basin),
10.2 miles east of Bismarck on State Route 84 (Sec. 35, R 19W, T 4S), Hot Spring
County, Arkansas. The crayfish were collected from shallow (8 to 16 cm) marginal
pool area choked with dense mats of filamentous algae and decaying organic
material on the south side of the road. The small springfed woodland stream
originates just north of the highway in a mixed Quercus-Pinus forest and flows
under the highway, through a pasture, and southward to join Prairie Bayou. The
stream bed consists of sandy clay interspersed with gravel, and grasses and sedges
flank its banks.

Disposition of types.—The holotype, allotype, and morphotype are deposited
in the National Museum of Natural History (Smithsonian Institution), nos. 177698,
177699, and 177700, respectively, as are the paratypes consisting of 2 dI, 11 oI,
12 2, 364346, 47j)2. See “‘Range and specimens examined’’ for restricted list of
types.

Size.—The largest specimen available is a first form male, from Sevier County,
that has a carapace length of 42.5 mm (postorbital carapace length 24.4 mm);

VOLUME 95, NUMBER 3 3) Il

corresponding lengths of the smallest first form male and largest female are 31.3
(25.8) mm and 39.6 (33.4) mm. Females carrying eggs or young are unknown.

Range and specimens examined.—All of the specimens of this crayfish of which
we are aware were collected in tributaries of the Ouachita and Red rivers in
southwestern Arkansas. In the following list of specimens examined, only those
lots marked by an asterisk constitute the type-series. CLARK COUNTY: (1)
small stream and roadside ditch 1.0 mi E of Amity on St Rte 84, 1 jd, 21 Apr
1973, J. E. Pugh, G. B. Hobbs, and HHH; (2) Wingfield Creek 0.5 mi E of St
Rte 53 on timber access road, 2 jd, 1j2, 8 Apr 1974, HWR; (3) Rest Haven
Cemetery about 4 mi W of Arkadelphia on St Rte 8, | dII, 28 Feb 1981, HWR;
(4)* creek 14.8 mi W of Bismarck on St Rte 84,7 dU, 4 2, 8jd, 1032, 13 Mar
1981, HWR. GRANT COUNTY: (5) creek 7.3 mi E of Poyen on St Rte 270, 2
2,154, 19 Mar 1980, HWR. HOT SPRING COUNTY: (6)* trib to Prairie Bayou
10.2 mi E of Bismarck on St Rte 84, 1 dI, 2 GI, 9jd,8j2, 17 Mar 1980, HWR;
22,655, 1032, 13 Mar 1981, HWR; 1 oI, 1j¢, 3 Apr 1981, HWR; (7)* roadside
ditch 6.4 mi W of Bismarck on St Rte 84, 2 dII, 3 2, ll jd, 1632, 17 Mar 1980,
HWR;; (8) trib to Point Cedar Creek 21 mi E of Point Cedar on St Rte 84, 1),
13 Mar 1981, HWR. NEVADA COUNTY: (9) De Ann Cemetery in Prescott, 2
ll, 2 2, 16jd, 16j)2, 28 Feb 1981, HWR. OUACHITA COUNTY: (10) trib to
Two Bayou between St Rtes 4 and 24, 1 dII, 136,452, 30 Mar 1975, S. O. Pelt.
PIKE COUNTY: (11) roadside ditch 2.0 mi E of Daisy on U.S. Hwy 70, 9jd, 6
j@, 21 Apr 1973, JEP, GBH, HHH. SEVIER COUNTY: (12) seepage area 5.0
mi NE of U.S. Hwy 59-71 on U.S. Hwy 70, 1 oI, 1 dU, 4 2, 1j3d,3j2, 20 Apr
1973, JEP, GBH, HHH.

Variations.—Most of the specimens at hand were collected in the Ouachita
River basin and exhibit a rather remarkable uniformity. Among the juveniles,
however, many of those with carapace lengths of 10 to 14 mm have rostra with
small marginal spines that appear to become reduced to angles with subsequent
molts, and, in most individuals, even the angles disappear completely before the
animal reaches sexual maturity. Among 15 juveniles from locality 11, those with
a carapace length greater than 14 mm do not have even a trace of the marginal
Spines on the rostrum or spines on the postorbital ridges. Some juveniles from
several of the localities lose both the marginal and postorbital spines by the time
they have attained a carapace length of 7 mm. In contrast, juveniles from locality
12 still have rostral and postorbital spines at a carapace length of 13 mm. The
juveniles from locality 9, Nevada County, have proportionately narrower areolae
than do those from the other localities. In only one of the adults (one from Sevier
County) is there even the slightest subangular curve marking the base of the
acumen that is so evident in the smallest juveniles. Also the fingers of the chela
of the single first form male from the same locality in Sevier County are slightly
more bowed than they are in the two first form specimens from Hot Spring
County. Inadequate series are available from the two river basins to ascertain
whether or not populations from them exhibit distinguishing features. Series of
adults from both drainage systems are much needed.

Relationships.—Procambarus (G.) parasimulans has its closest affinities with
P.(G.) simulans (Faxon, 1884:112) and P. (G.) curdi Reimer (1975:22). From P.
(G.) simulans it differs markedly in features of the cheliped: the chela is not only
more robust in P. (G.) parasimulans but it is also studded with more tubercles—

552 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Ratios of carapace lengths and areolar lengths and widths.

Ratios

Crayfishes il 7to 3t
P. (G.) parasimulans Range 30.0-34.5 37.8-42.3 5.1-9.0
(n = 34) Mean 32.5 + 1.2440 39.9 + 0.9619 6.6 + 0.9822
P. (G.) simulans (n = 15) Range 33.7-35.5 42.2-44.2 6.5-13.4
Mean 34.7 + 0.6488 43.4 + 0.5738 10.2 + 1.9779
P. (G.) curdi (n = 2) Range 36. 1—-37.5 44.0 14.6-16.0

* Areola length/Carapace length x 100. ** Areola length/Postorbital length x 100. + Areola length/
Areola width.

on the ventral surface distributed over the proximal half of the fingers, and on
the dorsal face of the dactyl, from base to midlength. Almost equally obvious is
the difference in the shape of the telson, which is narrower and more tapering in
the new species. The rostrum of the latter is also strikingly different in that the
margins are distinctly more gently convergent, almost all adult specimens lacking
even a suggestion of angles at the base of the acumen. Whereas the range of
variation in the relative length and width of the areola in the two overlap to some
extent, the areola of P. (G.) parasimulans is almost always shorter and broader
(see Table 2). The first pleopod of the first form male exhibits few differences:
the shoulder at the cephalic base of the terminal elements is less arched in P.
(G.) parasimulans, the flat surface of the caudal process lies more nearly parallel
to the transverse plane of the body, and the proximomesial spur is borne on the
caudal proximomesial angle of the appendage rather than being situated slightly
more laterally as in P. (G.) simulans simulans. Also the postannular sclerite of
the female is broadly rounded anteriorly in the new species rather than being
subtriangular.

The most obvious feature that distinguishes it from P. (G.) curdi is the areola
which is no more than 9 (as contrasted with 14 to 16) times as long as broad. The
postaxial surface of the ischium of the third maxilliped is much less hirsute, and
there are distinct differences in the first pleopod of the males of the two: the
central projection of P. (G.) parasimulans extends neither so far distally nor
laterally beyond the caudal process; in P. (G.) parasimulans the latter is obliquely
compressed, not mesiolaterally as it is in P. (G.) curdi.

This crayfish may be distinguished from the other species of the subgenus
Girardiella that have been reported to occur in Arkansas (see review by Bouchard
and Robison 1981) as follows: except for P. (G.) tulanei Penn (1953:163), which
may be recognized by the bearded mesial surface of the palm of the chela, the
areolae of these distant relatives [P. (G.) gracilis (Bundy, 1876:5), P. (G.) lib-
erorum Fitzpatrick (1978:533), and P. (G.) reimeri Hobbs, 1979:804)] are at least
16 times as long as wide, considerably narrower than that of P. (G.) parasimulans
of which no representative has been examined that has an areola greater than 9
times as long as wide.

Acknowledgments

We are grateful to G. B. Hobbs, Darryl D. Koym, Elaine E. Laird, Stephen
O. Pelt, and J. E. Pugh for their help in obtaining the specimens on which this

VOLUME 95, NUMBER 3 553

description is based and to J. F. Fitzpatrick, C. W. Hart, Jr., and H. H. Hobbs
III for their criticisms of the manuscript.

Literature Cited

Bouchard, Raymond W., and Henry W. Robison. 1981. An inventory of the decapod crustaceans
(crayfishes and shrimps) of Arkansas with a discussion of their habitats —Arkansas Academy
of Science Proceedings 34:22-30, 2 figures.

Bundy, W. F. 1876. In S. A. Forbes, List of Illinois Crustacea, with descriptions of new species.—
Bulletin of the Illinois Museum of Natural History 1:3, 4, 5, 24-25.

Faxon, Walter. 1884. Descriptions of new species of Cambarus, to which is added a synonymical
list of the known species of Cambarus and Astacus.—Proceedings of the American Academy
of Arts and Sciences 20:107-158.

Fitzpatrick, J. F., Jr. 1978. A new crawfish of the subgenus Girardiella, genus Procambarus from
northwest Arkansas (Decapoda, Cambaridae).—Proceedings of the Biological Society of Wash-
ington 9(2):533-538, 1 figure.

Hobbs, Horton H., Jr. 1979. A new crayfish from the Ouachita River basin in Arkansas (Decapoda:
Cambaridae).—Proceedings of the Biological Society of Washington 92(4):804-811, 1 figure.

Penn, George Henry, Jr. 1953. A new crawfish of the genus Procambarus from Louisiana and
Arkansas (Decapoda: Astacidae).—Journal of the Washington Academy of Sciences 43(5): 163—
166, 12 figures.

Reimer, Rollin D. 1975. Procambarus (Girardiella) curdi, anew crawfish from Arkansas, Oklahoma,
and Texas (Decapoda, Astacidae).—Tulane Studies in Zoology and Botany 19(1 and 2):22—25,
9 figures.

(HHH) Department of Invertebrate Zoology, Smithsonian Institution, Wash-
ington, D.C. 20560; (HWR) Department of Biology, Southern Arkansas Univer-
sity, Magnolia, Arkansas 71753.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 554-556

A NEW ULOBORUS LATREILLE SPECIES
FROM ARGENTINA 7
(ARACHNIDA: ARANEAE: ULOBORIDAE)

Brent D. Opell

Abstract.—The new species Uloborus elongatus is described and illustrated.

Uloborus elongatus is the eighth American uloborid species to be recognized
(Muma and Gertsch 1964; Opell 1979, 1981). Like U. glomosus (Walckenaer), U.
trilineatus Keyserling, and U. metae Opell, it has two stout tracheal trunks that
enter the cephalothorax, a long male conductor and convoluted epigynal ducts.
The remaining species, U. campestratus Simon, U. divisus Marx, U. eberhardi
Opell, and U. segregatus Gertsch, form a group characterized by four tracheal
trunks, a short conductor and simple epigynal ducts.

Uloborus elongatus, new species
Figs. 1-8

Types.—Female holotype and male paratype from Cataratas del Iquazu (25.40S,
54.27W) in the Argentinian province of Misiones, collected November 1963 by
Dr. Maria E. Galiano; deposited in Museo Argentino de Ciencias Naturales,
Buenos Aires.

Etymology.—The specific epithet refers to the slender carapace of this species.

Diagnosis.—Uloborus elongatus males and females are distinguished from those
of other species by white circles around anterior median eyes (AME’s) and by a
carapace which is 1.2 and 1.3, respectively (rather than 1.05), as long as wide
(Figs. 1, 5). Asin U. eberhardi, females have short epigynal lobes that are widely
separated at their bases (Fig. 4), but U. elongatus lacks the abdominal tubercles,
lateral spots, and club-shaped setae of this species. Males are distinguished by a
median apophysis bulb which in apical view (Fig. 8) is more nearly circular than
that of U. metae or U. trilineatus. Like the latter two species, U. elongatus has
a conductor which is four or five times as long as wide, but is distinguished from
both by a carapace length of only 1.05 mm and from the former by lacking lateral
abdominal black spots.

Description.—Female: Total length 3.28 mm, carapace length 1.12 mm, ster-
num length 0.66 mm. Carapace light tan with circular guanine deposits around
AME’s (Fig. 1). Posterior eyes ringed in black. AME diameter half that of others.
Sternum and legs tan without markings. First legs with evidence of worn tibial
setal brushes; femur 1.65 mm long without macrosetae, tibia with one central
dorsal and one distal retrolateral macroseta. Metatarsus IV 0.94 mm long; cal-
amistrum 0.46 mm long, composed of 24 setae. Ventral comb of leg IV 0.64 mm
long, composed of five metatarsal and 12 tarsal macrosetae. Abdomen without
tubercles or markings, with dense guanine deposits except in tan cardiac, lung
and spinneret regions (Fig. 2). Cribellum 0.34 mm wide. Weakly sclerotized epig-

VOLUME 95, NUMBER 3 555

Figs. 1-8. Uloborus elongatus: 1, Female carapace; 2, Lateral view of female abdomen; 3, Ventral
view of cleared epigynum; 4, Ventral view of epigynum; 5, Male carapace; 6, Prolateral view of male
first femur, patella, and tibia; 7, Retrolateral view of male left pedipalpus; 8, Apical view of male left
pedipalpus. S = spermatheca, FD = fertilization duct, CY = cymbium, E = embolus, C = conduc-
tor, MAB = median apophysis bulb, MAS = median apophysis spur.

556 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ynal lobes widely separated at bases (Fig. 4). Spherical spermathecae (Fig. 3)
visible through integument.

Male: Total length 2.48 mm, carapace length 1.04 mm, sternum length 0.58
mm. Carapace slightly darker than in female, with a more conspicuous median
stripe and less conspicuous white AME circles (Fig. 5). Femur I 1.40 mm long
with three prolateral and one retrolateral macrosetae; tibia I with seven prolateral,
11 dorsal and four retrolateral macrosetae (Fig. 6). Abdomen similar to female’s
except for a faint gray lateral stripe. Palpal femur with large retrolateral and small
prolateral proximal tubercles. Median apophysis bulb 0.14 mm wide with a shal-
low prolateral depression and a broad lobe adjacent to the long conductor tip
(Figs. 7, 8). Median apophysis spur pointed and more strongly bent than in other
species.

Distribution.—Known only from the type locality in northeastern Argentina.

Acknowledgments

I thank Dr. Maria E. Galiano for providing specimens and William Gorsica for
preparing carapace and abdomen illustrations.

Literature Cited

Muma, M. M., and W. J. Gertsch. 1964. The spider family Uloboridae in North America north of
Mexico.—American Museum Novitates, 2196: 1-43.

Opell, B. D. 1979. Revision of the genera and tropical American species of the spider family
Uloboridae.—Bulletin of the Museum of Comparative Zoology, Harvard 148(10):443-549.

—. 1981. New Central and South American Uloboridae (Arachnida, Araneae).—Bulletin of the
American Museum of Natural History 170(1):219-228.

Department of Biology, Virginia Polytechnic Institute, and State University,
Blacksburg, Virginia 24061.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 557-562

A NEW GENUS AND SPECIES OF POISON-DART FROG
(AMPHIBIA: DENDROBATIDAE) FROM THE
ANDES OF NORTHERN COLOMBIA

John D. Lynch and Pedro M. Ruiz-Carranza

Abstract.—A diminutive (3 adult females 18.6—19.3 mm SVL) dendrobatid frog
found in cloud forests on the Cordillera Central in Departamento Antioquia, Co-
lombia, is so unlike other dendrobatids that a new genus (Atopophrynus) is pro-
posed for the new species (A. syntomopus). Unlike other dendrobatids, A. syn-
tomopus has extensively webbed toes, has the innermost toe reduced in size and
fused to the second, and lacks ears.

In the course of preliminary sampling of the cloud forest frog fauna of the
Cordillera Central in northern Colombia, three specimens of a peculiar small frog
were collected along a small stream at the highest point on the road between
Sonson and Narino in southern Antioquia. Because the species presents a com-
bination of characteristics so markedly discordant with those previously reported
for dendrobatid frogs we here propose a new genus for it.

Atopophrynus, new genus

Diagnosis. —A dendrobatid frog distinguished from all other dendrobatids by
the absence of the disk and pad of the innermost toe of the hind foot and by
having that non-digitiform digit fused to the adjacent toe. Functional toes (II-V)
of hind foot extensively webbed and bearing large, rounded disks and pads. Ato-
pophrynus is also unique within the family in the absence of the tympanic annulus,
cavum tympanicum, and plectrum.

Type species.—Atopophrynus syntomopus Lynch & Ruiz.

Etymology.—From the Greek Atopos (=strange, out of place) + phrynus (toad);
gender masculine.

Atopophrynus syntomopus, new species
Figs. 1, 2

Holotype.—Instituto de Ciencias Naturales, Universidad Nacional de Colom-
bia (amphibian collection) 8611, one of three collected at the crest of the Cordi-
llera Central, 8 km by road E Sonson, Municipio Sons6n, Departamento Antio-
quia, Colombia, 2780 m, 13 June 1981, by Vincente Rueda.

Paratypes.—ICN 8612 and 8613 (cleared and stained skeleton), collected with
holotype.

Diagnosis.—same as for genus.

Description.—Head narrower than body, wider than long; snout broad, oval
(with a pointed tip) in dorsal view (Fig. 1), protruding in lateral profile, short;
nostrils directed anterolaterally, protuberant, canthus rostralis evident, rounded;
loreal region slightly concave, sloping abruptly to non-flared lips; interorbital

558 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Head of Atopophrynus syntomopus. Dorsal (ICN 8612) and lateral (ICN 8611) views. Line
equals 2 mm.

space broader than upper eyelid, flat; temporal region swollen (Fig. 1); no supra-
tympanic fold; no tympanic annulus; postrictal tubercle subconical; choanae round,
moderate-sized, lateral on palate but not concealed by palatal shelf of maxillary
arch; no vomerine odontophores or teeth; no teeth on premaxillae or maxillae;
tongue as long as wide, its posterior margin not indented, posterior % not ad-
herent to floor of mouth.

Skin of dorsum smooth, lacking folds but bearing a few scattered subconical
tubercles; skin of venter smooth; anal opening not extended in sheath; no enlarged
tubercles in vicinity of anus; no tubercles on limbs (ulnar and tarsal tubercles or
folds absent).

Palmar tubercle round, slightly larger than oval thenar tubercle; other tubercles
on hand very low, scarcely distinguishable; digits broad, basally webbed (Fig. 2);
fingers II-IV bearing obvious rounded pads; pads bearing disks on ventral sur-
faces (broader than long); thumb shorter than second finger, not bearing pad or
disk; ill-defined pair of scutes on tops of pads on fingers IJ-IV.

Outer metatarsal tubercle round, as large as, and more pungent (Fig. 2) than
inner metatarsal tubercle; latter flat, curved (kidney-shaped); no supernumerary
plantar tubercles; foot appearing to have only four toes (Fig. 2); toes II—V bearing
low subarticular tubercles, nearly fully webbed (as a fringe on distal portion of
toe IV), bearing broad disks on expanded, apically round digital pads; inner
margin of toe II bearing fleshy flange along *% of its length; although lacking pad,
disk, and subarticular tubercle, this flange containing first toe; hind limbs short,
heels of flexed hind legs (when held at right angles to sagittal plane) not meeting.

In preservative, gray above with reddish-brown markings (interorbital bar,
scapular and sacral chevrons, blotch on lower back, limb bars, slanted bars on
flanks) and flecked with white (most intense on lateral surfaces); ventral surfaces
cream, blotched with white and flecked with brown; pair of brown blotches on
breast; inner digits pale cream.

In life dorsum red with green markings overlain with white flecks; white flecks

VOLUME 95, NUMBER 3 sy)

“

Fig. 2. Palmar views (A) ICN 8612, (B) ICN 8613, and plantar views (C) ICN 8611, (D) ICN 8613,
of acropodia of Atopophrynus syntomopus. Line equals 2 mm.

560 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

most dense on flanks; undersides pale olive with blue-white spots edged with
brown; digital tips yellow; iris bright copper; posterior surfaces of thighs olive-
green.

ICN 8613 was prepared as an Alizarin cleared and stained skeleton. Surface
color of muscles cream; nasal bones very small, situated well lateral on snout,
widely separated from all other dermal bones; frontoparietals not ossified medi-
ally, exposing long narrow fontanelle; maxillary arch complete, edentate; alary
processes of premaxillae directed anterodorsally; pars facialis of maxilla low;
bones of maxillary arch very thin aside from broad palatine processes of pre-
maxillae and palatal shelf of adjacent portions of maxillae; otic ramus of squa-
mosal longer than ascending ramus, zygomatic ramus very short (not as long as
deep); vomers C-shaped bones bordering inner margins of choanae; thin palatine
bones present, extending from sphenethmoid to maxillae; median ramus of pter-
ygoid obsolete; anterior ramus of pterygoid long and in broad contact with maxilla
but not reaching palatine.

Neural arches of eight procoelous presacral vertebrae not imbricate; atlantal
cotyles widely separated; sacrococcygeal articulation bicondylar; phalangeal for-
mulae 2-2-3-3 and 2-2-3-4-3; except for innermost digit, terminal phalanges broad-
ly T-shaped (Fig. 2); pectoral girdle firmisternal; clavicles thin, not reaching mid-
line; coracoids, especially medial portions, large; no omosternum detected; sternum
essentially undetectable (not so large as that illustrated for Dendrobates histri-
onicus by Silverstone 1975:6, fig. 2b).

Measurements in mm.—Measurements are for ICN 8611 (holotype), 8612, and
8613, respectively. SVL 19.3, 19.3, 18.6; shank 8.2, 8.4, 7.9; HW 7.0, 6.2, 6.1;
head length (estimated) 5.7, 5.3, 5.9; chord of head length 6.6, 6.2, 5.7; upper
eyelid width 1.6, 1.5, 1.6; IOD 2.3, 2.2, 2.2; eye length 2.5, 2.4, 2.2; E-N I-5;
1.4, 1.6.

Natural history.—The three frogs captured were the only frogs seen along the
small (ca. 0.5 m wide), deep (2.5 m), rock-walled, densely shaded ravine by day.
At night several species of Eleutherodactylus, Hyla larinopygion, Hylopsis buck-
leyi, and Rhamphophryne sp. were found in the vicinity of the stream. Searching
the stream by day failed to reveal any other specimens, but those that were found
were disturbed from crevices in the rock forming the bottom and walls of the
stream. The frogs were quite active and when jumping behaved much like Co-
lostethus (e.g., C. abditaurantiacus, C. palmatus, and C. vergeli) in first hopping
onto rock faces under tiny waterfalls and, if pursued further, leaving the imme-
diate vicinity of discovery by hopping into the water and being quickly washed
downstream. Each specimen is an adult female with strongly convoluted oviducts
and large ovarian eggs. ICN 8613 contains six large ova (1.8—2.0 mm in diameter)
in each ovary (as well as numerous much smaller ova). No tadpoles of any kind
were found in the stream (the section searched lacked any pools).

Etymology.—Syn + tomos (=abridged) + pous (=foot), Greek.

Discussion

A persistent concern as to the generic limits within the biologically and phar-
maceutically interesting poison-dart frog family Dendrobatidae has plagued her-

VOLUME 95, NUMBER 3 561

petologists for much of this century. The three currently recognized genera (Co-
lostethus, Dendrobates, and Phyllobates) present a mosaic distribution of attributes
commonly used to discriminate between frog genera overlain by an as yet incom-
plete biochemical survey. Edwards (1974), Savage (1968), and Silverstone (1975,
1976) divided the family on the basis of negative attributes (1.e., plesiomorphic
features and their absences in subsets of dendrobatids). The use, by these authors,
of the presence/absence of “‘bright’’ coloration (not defined) continued a tradition
which was overthrown by Myers et al. (1978) who delimited Phyllobates and the
suprageneric unit consisting of Dendrobates + Phyllobates on the bases of bio-
chemical synapomorphies. As noted by Myers et al. (1978), the resulting classi-
fication is not entirely satisfactory because there are no non-biochemical attri-
butes corroborating it. However, there are no apparent attributes distributed in
such a fashion as to serve as falsifiers of that classificatory hypothesis, and it
stands as the least refuted hypothesis of relationships.

Our ability to place Atopophrynus into the classificatory hypothesis of the
Dendrobatidae is impaired by the apparent lack of synapomorphies for Coloste-
thus (cf. Myers et al. 1978) and by the absence of data on reproductive behavior
in Atopophrynus. Excluding Atopophrynus and two species provisionally referred
to Colostethus by Lynch (1982), all other dendrobatids have an inner tarsal fold
or tubercle. The habit of transporting the tadpoles on the parent’s back, so well-
known in Dendrobates and Phyllobates (Silverstone 1975, 1976) and most Co-
lostethus (Edwards 1974), is not known in Atopophrynus nor in the two species
of Colostethus described by Lynch (1982). Each of these features is one poten-
tially fragmenting Colostethus. Neither the peculiar modification of the hind foot
of Atopophrynus nor its loss of the ear is presaged elsewhere in the family. The
absence of dentition in Atopophrynus is an apomorphy of low value (because it
postulates the loss of an attribute; see Hecht and Edwards 1976); we do not take
it as sufficient evidence that Atopophrynus is closely related to other edentulous
dendrobatids.

Acknowledgments

Fieldwork in Colombia was partially supported by funds from the Universidad
Nacional de Colombia and the University of Nebraska Research Council. For
aid in the field we thank Pablo Bernal and Vincente Rueda.

Literature Cited

Edwards, S.R. 1974. A phenetic analysis of the genus Colostethus (Anura: Dendrobatidae).—Ph.D.
dissertation, University of Kansas, 419 pp.

Hecht, M. K., and J. L. Edwards. 1976. The determination of parallel or monophyletic relationships:
the proteid salamanders—a test case.—American Naturalist 110:653-677.

Lynch, J. D. 1982. Two new species of poison-dart frogs (Colostethus) from Colombia.—Herpe-
tologica 38 :366-374.

Myers, C. W., J. W. Daly, and B. Malkin. 1978. A dangerously toxic new frog (Phyllobates) used
by Embera indians of western Colombia with discussion of blowgun fabrication and dart poi-
soning.—American Museum of Natural History Bulletin 161:307—366.

Savage, J. M. 1968. The dendrobatid frogs of Central America.—Copeia 1968:745-776.

562 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Silverstone, P. A. 1975. A revision of the poison-arrow frogs of the genus Dendrobates Wagler.
Natural History Museum of Los Angeles County, Science Bulletin 21:1—55.

1976. A revision of the poison-arrow frogs of the genus Phyllobates Bibron in Sagra (family
Dendrobatidae).—Natural History Museum of Los Angeles County, Science Bulietin 27:1—S3.

(JDL) School of Life Sciences, The University of Nebraska, Lincoln, Nebraska
68588; (PMR-C) Instituto de Ciencias Naturales, Universidad Nacional de Co-
lombia, Apartado Aereo 7495, Bogota, D.E., Colombia.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 563-566

A NOTE ON THE GENITALIA OF POTAMOTHARIX
HAMMONIENSITS (OLIGOCHAETA: TUBIFICIDAE)

H. R. Baker

Abstract.—Potamothrix hammoniensis (Michaelsen, 1901) is shown to possess
a thickened internal penial basement membrane that is known elsewhere only in
Haber Holmquist, 1978. This character is present in the syntype material of P.
hammoniensis and in English and Italian specimens. A description of the syntypes
of P. hammoniensis is given.

Examination of syntype material of Potamothrix hammoniensis (Michaelsen,
1901) has revealed that while an external thickened cuticular penis sheath is
absent, an internal thickened penial basement membrane previously unreported
is present. To date, this character is known elsewhere only from the genus Haber
(Holmquist 1978, 1979). As P. hammoniensis is a very common European fresh-
water oligochaete, a complete description is desirable to ensure the stability of
this taxon. Although this paper is not intended to be a formal redescription of P.
hammoniensis, a complete description is given of the syntype material in order
to supplement the original description. For a full synonymy of P. hammoniensis
see Brinkhurst (1971).

Methods and Materials

Three syntypes of Ilyodrilus hammoniensis Michaelsen, 1901 were loaned to
me courtesy of Dr. E. G. Easton, Annelida Section, British Museum (Natural
History) (BMNH). Italian specimens of P. hammoniensis from L. DiGarda and
L. d’Endine were collected by K. Coates and Dr. G. Bonomi; English specimens
from Shropshire were collected by Ms. S. Cowley. The anterior part of one
syntype was sectioned by the Laboratory of Analytical Systematics, Royal On-
tario Museum, Toronto, Ontario. The second syntype, the posterior part of the
sectioned specimen, and the Italian and English specimens were stained in para-
carmine, acid-differentiated, dehydrated through an ethanol-xylene series and
mounted in Canada balsam. The third syntype was retained in alcohol.

Systematic Description
Potamothrix hammoniensis (Michaelsen, 1901)
Figs. 1, 2

Ilyodrilus hammoniensis Michaelsen, 1901

Description (syntypes).—Length 12.4 mm and 15 mm, 75 and 71 segments
respectively, width at clitellum (in unsectioned, slightly compressed, whole
mounted specimen) 950 um, 500 um in sectioned specimen. Prostomium conical
with rounded tip, about as long as broad at peristomium. Clitellum over XI—XII,
absent ventrally in XI between male pores. Anterior dorsal setal bundles with 2—
4 pectinate setae and 1-4 straight hair setae; pectinate setae with upper tooth
longer and thinner than lower (2-3 small intermediate teeth often present), from

564 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

300 pm

Fig. 1. Potamothrix hammoniensis, lateral view of genitalia in segments X and XI (drawn from
syntype). a, atrium; bm, basement membrane; m, muscle layer; pa, penial apparatus; pr, prostate;
ps, penial sac; sf, sperm funnel; sp, spermatheca; ss, spermathecal seta; vd, vas deferens.

125-140 wm long, 4-6 um thick; hair setae from 330-400 um long, approximately
2.5 ym thick. Posterior dorsal bundles with 1-2 bifid and 1-2 hair setae, bifid
setae may have some small intermediate teeth; both bifid and hair setae decrease
slightly in size posteriorly. Anterior ventral bundles with 3-5 bifid setae, posterior
ventral bundles with 2—4 bifid setae; upper tooth longer and thinner than lower.
Ventral setae of X modified as spermathecal setae (ss; Fig. 1, 2A); each bundle
contains one large, thick (240 um long; maximally 11 um wide entally, 13.7 wm
wide ectally) seta with distal hollow, gutter-shaped tip; tip of seta sharply pointed.
Dorsal somatic setae of X present, ventral somatic setae of XI absent. Sper-
mathecal and male pores median in X and XI respectively, spermathecal pores
dorsolateral to spermathecal setae, in or slightly ventral to lateral line; male pores
in ventral setal line. Chloragogen cells thick over gut in VI and VII, thinner
posteriorly. Anus subterminal.

Male system (all structures paired) (Fig. 1, 2B): sperm funnel (sf) large, asym-
metrical, ventral lip larger. Vas deferens (vd) short (approximately 30-40 um
long) and wide (23-26 um), joining atrium apically. Atrium (a) 44 wm wide at
junction of vas deferens, 98 wm wide at junction of prostate gland (pr), muscle
layer (m) 6.5 um thick in region of prostate gland junction; prostate gland joins
atrium approximately 220 ~m from sperm funnel. Atrium very long and tubular
(1900 um long to apical end of penial apparatus, 43-187 wm wide) with wide
lumen; joining penial apparatus apically. The penial apparatus (Fig. 2B) consists
of a muscular bulb the ectal part of which forms the main part of the penis; the
penis is contained in a penial sac (ps) and lacks an external thickened cuticular

VOLUME 95, NUMBER 3 565

100 pm

Fig. 2. Potamothrix hammoniensis: A, Spermathecal seta from syntype; B, Section of penial
apparatus showing entrance of atrium (from L. d’Endine specimen), notation as in Fig. 1. Tissue over
basement membrane represented by stippling.

Sheath. Atrial tissue runs through the muscular bulb to join the very thin layer
of epithelial tissue covering the penis; penial epithelial tissue continuous with
penial sac epithelial tissue. Basement membrane (bm) of penial epithelial tissue
thickened over protrusible part of penis except for the most ectal tip of penis.

Spermathecae (sp; Fig. 1) with distinct ducts and large saccular ampullae; ducts
bulbous basally (55-60 um wide) narrowing to 25-30 wm above bulbous bases,
gradually widening to join ampullae, duct length 170-180 m. Thin muscle layer
(2.5—3.0 um thick) over ducts of spermathecae, muscle layer very thin or absent
Over ampullae. Sperm trap absent. Sperm as spermatozeugmata.

Remarks.—The degree of pectination of the dorsal setae tends to become re-
duced posteriorly so that some of the most posterior dorsal bundles contain bifid
setae with no sign of pectination.

Material examined.—BMNH 1903. 4.28.10-14, 3 syntypes of P. hammonien-
sis; Baker collection: 20 whole-mounted specimens and 2 sectioned specimens
from L. di Garda and L. d’Endine, Italy, 2 whole-mounted specimens from
Shropshire, England; Cowley collection: 9 whole-mounted specimens from
Shropshire, England.

Discussion

Potamothrix hammoniensis 1s the only member of Potamothrix that is known
to possess a thickened penial basement membrane. This thickened membrane is
known elsewhere only in Haber (Holmquist 1978, 1979). Although this feature
appears to be rare within the Tubificidae, it is not suggested that P. hammoniensis
is related to the members of Haber (recently expanded by Brinkhurst 1981). In

566 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Potamothrix the vasa deferentia are very short and modified penial setae are
absent (Brinkhurst 1971), whereas Haber has long vasa deferentia (composed of
distinct thin and thick parts) and modified penial setae (Holmquist 1978).

The Italian and English specimens conform closely to the description of the
syntypes. However, most of the specimens bear I—2 normal ventral somatic setae
in XI immediately posterior to the male pore. These setae are absent in the
syntypes. The muscular bulb of the penial apparatus is usually also slightly larger
than those of the syntypes. These differences are regarded here as being minor.

Michaelsen (1901:67) described the penis as ““Penis weich, bei vallstandiger
Ausstreckung in der Mitte knopformig angeschwollen, am freien Ende zapfen-
formig.’’ He apparently did not notice the thickened basement membrane of the
penis. The thickening of the epithelial basement membrane in the penis may act
as a penial stiffening device in much the same manner as an external cuticular
penis sheath. This thickened basement membrane also provides an insertion point
for the muscles of the penial bulb.

Acknowledgments

My thanks to Dr. R. O. Brinkhurst for providing working facilities and con-
structive criticism as it was needed. I thank Ms. S. Cowley, Ms. K. Coates and
Dr. G. Bonomi for allowing me to examine material which they collected; and
Mr. Cary Gilmour (Royal Ontario Museum) for sectioning the specimen of P.
hammoniensis. | am indebted to Dr. E. G. Easton for the loan of syntype material
of P. hammoniensis and for his patience concerning the loan. This manuscript
was typed by M. Stone. Financial support was provided by a Natural Sciences
and Engineering Research Council of Canada post-graduate award.

Literature Cited

Brinkhurst, R. O. 1971. Jn Brinkhurst, R. O., and B. G. M. Jamieson. Aquatic Oligochaeta of the
World. Oliver and Boyd, Edinburgh (Part 2. Systematics. 8. Family Tubificidae. pp. 444-625).

—. 1981. A contribution to the taxonomy of the Tubificinae (Oligochaeta: Tubificidae).—Pro-
ceedings of the Biological Society of Washington 94(4): 1048-1067.

Holmquist, C. 1978. Revision of the genus Peloscolex (Oligochaeta, Tubificidae). 1. Morphological

and anatomical scrutiny; with discussion on the generic level.—Zoologica Scripta 7:187—208.

. 1979. Revision of the genus Peloscolex (Oligochaeta, Tubificidae). 2. Scrutiny of the species.—

Zoologica Scripta 8:37—-60.

Michaelsen, W. 1901. Neue Tubificiden des Niederelbgebietes.—Verhandlungen des Naturwissen-
schaftlichen Vereins Hamburg 3(8):66—70.

Department of Biology, University of Victoria, P.O. Box 1700, Victoria, British
Columbia, Canada V8W 2Y2; Research Associate, British Columbia Provincial
Museum, Victoria, British Columbia, Canada.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 567-570

FRESHWATER TRICLADS (TURBELLARIA) OF NORTH
AMERICA. XIV. POLYCELIS MONTICOLA,
NEW SPECIES, FROM THE SIERRA
NEVADA RANGE IN CALIFORNIA

Roman Kenk and Anne M. Hampton

Abstract.—A new species of Polycelis, P. monticola, is described. The prin-
cipal distinguishing characters of the species are in the anatomyof the copulatory
apparatus. The species has a blunt penis papilla and the ejaculatory duct opens
on the ventral side of the papilla instead of at its tip.

In the course of a study on the distribution and ecology of freshwater triclads
or planarians in the Lake Tahoe Basin, California and Nevada, one of us (AMH)
found representatives of the genus Polycelis to be the most common planarians
inhabiting springs and creeks of the basin. As Polycelis sierrensis Kenk was
known to be present in the area (Kenk 1973:11), it was natural to assume that all
the specimens observed belonged to that species. Upon closer examination, how-
ever, and after studying the anatomy of sexually mature individuals, it was dis-
covered that some of them belonged to a new species.

Polycelis monticola, new species

Type-material.—All type-specimens have been deposited in the National Mu-
seum of Natural History, Smithsonian Institution, Washington, D.C.

Holotype: whole mount of anterior end and sagittal sections of posterior part
on 7 slides (USNM 71762). Paratypes: whole mount and sagittal sections of 5
specimens, on 44 slides (USNM 71763-71767).

External features (Fig. 1).—In general external appearance the species con-
forms with the other two American species of Polycelis, P. coronata and P.
sierrensis. Fully grown specimens are up to 20 mm long and 2.5 mm wide. The
anterior end is truncate, with a convex frontal margin and a pair of rather pointed
auricles extending anterolaterally. Behind the auricles, the lateral body margins
gradually widen, soon reach their greatest width, run parallel for some distance,
then converge again behind the pharyngeal region, to meet at the bluntly pointed
posterior end. The color of the animal is a variable shade of brown, somewhat
darker on the dorsal surface than ventrally. The numerous small eyes are arranged
along the margins of the head and the anterior fourth of the prepharyngeal region.
They are rather widely scattered on the head and may be absent or scarce on the
bases of the auricles (Fig. 2A). In the genus Polycelis, the individual eyes or
ocelli lack the unpigmented spots located above each ocellus, such as are found
in Dugesia or Phagocata. In darkly colored specimens it is therefore often dif-
ficult to see the small pigment cups of the individual ocelli through the colored
tissues covering them. The pharynx is rather long, measuring about one-fourth
the body length; its root is situated at the middle of the body.

Anatomy.—The distinguishing characters of the species are in the morphology

568 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Polycelis monticola, outline drawing of living specimen, indicating the location of eyes
and pharynx.

of the reproductive system. The gonads and gonoducts are very similar to those
of other species of the genus. The ovaries are situated on the medial side of the
ventral nerve cords at about the level of the fifth lateral branches of the anterior
intestinal trunk. The numerous testicles are arranged in a pair of longitudinal
bands beginning immediately behind the ovaries and extending posteriorly to the
root of the pharynx. They are predominantly ventral, located chiefly medially to
the ventral nerve cords, and connect with the pair of thin anterior sperm ducts
or vasa deferentia that run along the medial side of the cords. In the region of
the pharynx, the sperm ducts are widened, forming the false seminal vesicles or
spermiductal vesicles that proceed posteriorly to their entrance into the penis
bulb.

In the copulatory apparatus (Fig. 3), the gonopore (gp) leads into a short canal
that continues dorsally into the bursal duct (bd) and on the right side into the
male atrium (am). At the junction of these canals there is no widened cavity that
could be termed a common atrium. The musculature of the male atrium is of the
common type, not excessively developed, and consists of a layer of circular fibers
adjoining the epithelial lining, followed by a layer of longitudinal muscles. The
penis consists of an anterior bulb of moderate size and a plug-shaped or bluntly
conical papilla projecting into the male atrium. The bulb contains a large cavity
of irregular outline, the seminal vesicle (vs), from which a narrower canal, the
ejaculatory duct (de), proceeds to open ventrally to the tip of the papilla. Nu-
merous gland ducts (g/), with a granular, slightly eosinophilic secretion, enter the

Fig. 2. Photographs of anterior ends of preserved specimens, x20: A, Polycelis monticola from
Tahoe Paradise; B, P. sierrensis from spring near Sagehen Creek Biological Station, Nevada County,
California.

VOLUME 95, NUMBER 3 569

|
|
| I | | | I

0.5mm b vd gl vs de bd amodc gp
Fig. 3. Polycelis monticola, semidiagrammatic view of copulatory apparatus in sagittal section.

am, male atrium; b, copulatory bursa; bd, bursal duct; de, ejaculatory duct; gl, gland ducts; gp,
gonopore; odc, common oviduct; vd, vas deferens; vs, seminal vesicle.

penis bulb from the surrounding mesenchyme and apparently open into both the
penial lumen and the outer surface of the penis papilla. The two sperm ducts (vd)
enter the penis bulb laterally and open separately into the seminal vesicle. The
oviducts or ovovitelline ducts ascend dorsally in the region of the copulatory
complex, approach the midline, and unite above the male atrium to form the
common oviduct (odc), that curves posteriorly and opens into the end part of the
male atrium close to its junction with the bursal duct.

The copulatory bursa (b) is a large sac, rather elongate in well extended spec-
imens. Its duct, the bursal stalk (bd), proceeds posteriorly above the penis and
male atrium, slightly left of the midline, then curves ventrally toward the gono-
pore. The duct is surrounded by a strong muscle coat consisting of intermingled
circular and longitudinal fibers.

None of the epithelia of the copulatory complex is infranucleate.

Distribution and ecology.—Sexually mature, and therefore reliably identifiable,
specimens could be examined only from a small spring at Tahoe Paradise, El
Dorado County, California. The spring is located about 250 m east of State Road
89, 2 km south of its junction with U.S. Highway 50 (U.S. Geological Survey,
Primary base series map Echo Lake, California, Forest Service No. 535-4C, NE
quarter of NW quarter of Section 5, Township 11 N, Range 18 E). Many indi-
viduals of Polycelis, however, were found in other springs and creeks of the area,
which had their reproductive systems developed only in the anlage stage, not
sufficiently differentiated to separate them from other species of the genus.

Another, though less dependable character, the scattered arrangement of the
eyes on the head region of P. monticola, distinguishes it to some extent from the
sympatric P. sierrensis. This character indicates a wider distribution of the new
species in the Lake Tahoe area:

El Dorado County, California: Two more springs at Tahoe Paradise; spring
and a small creek at Benwood Meadows; Heavenly Valley Creek; Cold Creek.

Douglas County, Nevada: Springs on Foothill Road, Gardnerville.

The reproduction of P. monticola appears to be both sexual and asexual. Spec-
imens with regenerating anterior or posterior ends, indicating previous fissions,
were Observed at all seasons. Individuals with sexual structures were likewise

570 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

seen at all seasons, but no egg capsules or cocoons were found at any time despite
diligent searching.

Polycelis monticola was active in the Tahoe Paradise spring all year-round and
was collected at water temperatures varying between 1.1° and 16.0°C, pH 7.0.
However, the species is very sensitive to temperature fluctuations. When re-
moved from the spring and taken to the laboratory, animals often disintegrated.
This condition was exacerbated during the warmer months, presumably when
they were at the upper limits of their temperature tolerance. Repeated attempts
to send live specimens from California to Washington, D.C., in vacuum-insulated
bottles shipped by air mail special delivery, were successful only a few times.

Taxonomic position.—Polycelis monticola is closely related to P. coronata
(Girard), from which it differs chiefly in the anatomy of the male copulatory organ.
In P. coronata the penis papilla is conical and the opening of the ejaculatory duct
is at the rather pointed tip of the papilla. Polycelis monticola has a plug-shaped
or bluntly pointed papilla and the opening of the ejaculatory duct is ventral to
the tip. In an earlier paper (Kenk 1972:25-27), three subspecies of P. coronata
were distinguished. Two of them were separated only by the dfferentiation of
the bursal duct, which in P. c. coronata is divided into an anterior glandular and
a posterior muscular section, while in P. c. borealis Kenk it is muscular through-
out its length. Recent investigations have shown that various intergrades exist
between these two conditions. It is therefore best to abolish this distinction be-
tween the two subspecies and to consider P. c. borealis to be a synonym of P.
c. coronata. The third subspecies, P. c. brevipenis Kenk, distinguished by an
elongated penis bulb and a short penis papilla, seems to be a valid taxon.

Another species of Polycelis, P. sierrensis Kenk (1973), occurring also in the
Lake Tahoe area, differs from P. monticola by having the male atrium surrounded
by a very thick muscle coat, which places it in the subgenus Seidlia. In life it
can hardly be separated from P. monticola except that the band of eyes on the
head is more marginal in P. sierrensis, while in P. monticola the eyes are scat-
tered on the surface of the head (Fig. 2) and the auricles are more acutely pointed.

Etymology.—The name of the new species, monticola (Latin, inhabitant of
mountains), refers to its occurrence at high altitudes.

Acknowledgments

Thanks are due to Dr. John C. Harshbarger, in whose laboratory the photo-
graphs were prepared, and to Dr. Marian H. Pettibone, who kindly reviewed the
manuscript.

Literature Cited

Kenk, R. 1972. Freshwater planarians (Turbellaria) of North America.—Biota of Freshwater Eco-
systems, Identification Manual, No. 1. ix + 81 pp. Washington, D.C.: Environmental Protec-
tion Agency.

———. 1973. Freshwater triclads (Turbellaria) of North America. V. The genus Polycelis.—Smith-
sonian Contributions to Zoology 135. iii + 15 pp.

(RK) Department of Invertebrate Zoology, Smithsonian Institution, Washing-
ton, D.C. 20560; (AMH) Biological Research Associates, Box 11627, Tahoe Par-
adise, California 95708.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 571-574

REDESCRIPTION OF THE MAJOR SPINES
OF POLYDORA LIGNI WEBSTER
(POLYCHAETA: SPIONIDAE)

Jerry D. Kudenov

Abstract.—Polydora ligni Webster (Polychaeta: Spionidae) exhibits a generally
unreported accessory flange on the major spines of setiger 5 resembling that seen
in P. websteri Hartman, and P. haswelli Blake and Kudenov. This diagnostically
important flange was previously mentioned by Foster (1971) but has otherwise
not been commented on by other workers. Polydora ligni is compared with re-
lated polydorids in light of this morphological evidence.

Polydora ligni Webster (Polychaeta: Spionidae) was originally described as
having a single subdistal accessory tooth on the major spines of setiger 5 (Webster
1879:119). Numerous specimens from southern California and other locations in
North America were found to bear a subdistal, laterally produced flange of the
type seen in P. websteri Hartman, and also P. haswelli Blake and Kudenov. This
led to a re-examination of Webster’s types. Although this accessory flange was
originally mentioned by Foster (1971:22-24, fig. 19), it was inadequately described
and illustrated, and its significance was not fully discussed. As a result, this
systematically important feature has since been overlooked (for example, see
Blake 1971:5—6, 1975:210; Blake and Kudenov 1978:248; Light 1978: 175—177; Mi-
chaelis 1978; Rice and Simon 1980).

The purpose of this paper is to provide additional illustrations of the accessory
flange of the major spines of P. ligni, and to compare this species with closely
related forms in view of this character.

Polydora ligni Webster
Fig. |

Polydora ligni Webster, 1879:119, pl. 5, figs. 45-47.—Blake, 71:5-6, fig. 102 (syn-
onymy).—Foster, 1971:22—24, figs. 8-21 (synonymy).—Blake and Kudenov,
1978:248, fig. 43h-j.—Light, 1978:175-178, fig. 176 (synonymy).

Material.—NEW JERSEY: Great Egg Harbor, June-July 1878, H. E. Webster
coll., 15 types plus 3 slides (USNM 376). VIRGINIA: York River, from oyster
clusters, 11 June 1940, P. S. Galtsoff coll., 4 specimens (AHF N1510). NORTH
CAROLINA: Morehead City, at causeway, in water-soaked wood, 11 July 1940,
O. Hartman coll., 15+ specimens (AHF N1556); Beaufort, outer side of town
marshes, on soaked wood, in soft mud, debris and in old Teredo burrows, 21
June 1940, O. Hartman coll., 10+ specimens (AHF N1557). TEXAS: Point Penes-
cal, Laguna Madre, in galleries of serpulid tubes, 11 March 1950, W. A. Price
coll., 30 specimens (AHF N5648). CALIFORNIA: Santa Monica, Marina del
Rey, Station 10A, soft mud, 3 m, 13 April 1979, D. Soule and M. Oguri colls.,

572 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Polydora ligni (type, USNM 376). Major spines of setiger 5, dorsal view.

50+ specimens (AHF 000365-0); San Francisco Bay, San Pablo Channel, Station
17, August 1951, 50+ specimens (AHF N10397); Marin County, Drakes Estero,
11 May 1933, O. Hartman coll., 8 specimens (AHF N3447). BRITISH COLUM-
BIA (CANADA): Vancouver Island, Kyc Bay, E. Berkeley coll., 3 specimens
(AHF N1420).

Emended diagnosis.—Prostomium anteriorly bifurcate, with occipital cirrus;
notosetae lacking on setiger 1; setiger 5 typically lacking dorsal superior and
ventral inferior setae; major spines of setiger 5 smooth, with small lateral tooth
and accessory flange (Fig. 1) in unworn examples; companion setae distally bi-
furcate, feather-like; bidentate hooded hooks from setiger 7 unaccompanied by
capillaries, bearing distinct manubrium and with main fang forming acute angle
with shaft; pygidium flaring, cup-like, notched dorsally, not divided into lobes.

Remarks.—The above diagnosis basically follows those presented by Foster
(1971:22) and Light (1978:169). Foster’s mention of, but subsequent lack of dis-
cussion about the accessory flange is confusing, since this structure represents
both a significant systematic character and a modification of the concept of this
species. The presence of the flange is most easily detected in unworn spines (Fig.
1); it is generally absent in older, worn ones (as indicated by Foster). In addition,
the flange may be obscured by the companion setae (which are omitted from Fig.
1 for clarity). It is difficult to see the flange in ventral or lateral views, but it is
easily seen in the dorsal aspect under oil immersion. The fact that it has been
missed so consistently in this well-studied species is striking.

Polydora ligni further exhibits variability in the presence of superior and in-
ferior fascicles of capillary setae on setiger 5. Typically, both of these fascicles
are absent (Webster 1879:119; Sdderstr6m 1920:265; Blake 1971:5—6; Light 1978:

VOLUME 95, NUMBER 3 SS)

176-177). However, Rice and Simon (1980) encountered a high proportion of
individuals having both dorsal and ventral fascicles in founding populations at
Ballast Point, Florida, and Blake and Kudenov (1978:258) noted that a ventral
inferior fascicle is invariably present in Australian specimens of P. ligni. It is
therefore suggested that the presence or absence of these setal fascicles in such
eurytopic species as P. ligni is to be expected, especially in founding populations
(see Rice and Simon 1980). However, the Australian specimens identified as P.
ligni by Blake and Kudenov (1978:258) invariably possessed ventral inferior fas-
cicles on setiger 5, and probably represent an undescribed species.

Polydora ligni belongs to the basal P. ciliata-P. websteri major complex, to
which also belongs the P. alloporis-P. wobberi-P. cavitensis subgroup, since the
hooded hooks have a manubrium and with the main fang and shaft forming an
acute angle. Within this scheme, P. ligni is closely allied with P. latispinosa Blake
and Kudenov, which lacks dorsal superior fascicles of capillaries on setiger 5,
and has distally hastate companion setae. It is also related to other members of
the P. ciliata-P. websteri complex in having accessory structures on the major
spines of setiger 5. For example, an accessory tooth is present in both P. ciliata
Johnston and P. limicola Annenkova; two flanges are present in P. haswelli; and
a flange and a tooth are present in P. websteri Hartman. Light (personal com-
munication) has seen many specimens of P. websteri, especially in the Gulf of
California, and living commensally in Allopora venusta off California, in which
the flange and accessory tooth were not connected, much as that exhibited by P.
ligni.

It is of interest to note that Light (1978:177) compares P. ligni to two other
superficially similar species, namely P. nuchalis Woodwick and P. cirrosa Rioja.
These three species all have either occipital or nuchal cirri. However, the pres-
ence or absence of such structures is much less conservative than the structure
of the hooded hooks and major spines (see Rice and Simon 1980; Kudenov 1982).
The hooded hooks of P. nuchalis lack a manubrium and the main fang and shaft form
an obtuse angle. It is therefore clear that P. nuchalis is quite unrelated to P. ligni.
Polydora nuchalis is best considered with the P. socialis-P. flava-P. giardi major
complex to which a related species, P. tentaculata Blake and Kudenov, is already
assigned. Polydora cirrosa, on the other hand, possesses hooded hooks similar
to those of P. ligni, and appears most closely allied with the P. alloporis-P.
wobberi-P. cavitensis subgroup in view of the modified major spines, and the P.
ciliata kind of hooded hooks.

Acknowledgments

I am indebted to M. L. Jones, National Museum of Natural History, for lending
type-material of P. ligni; to R. Osborne, Allan Hancock Foundation, University
of Southern California, for drawing my attention to this problem; and to W. J.
Light, Department of Biology, University of Wisconsin, and K. Fauchald, Na-
tional Museum of Natural History, for critically reviewing this paper.

Literature Cited

Blake, J. A. 1971. Revision of the genus Polydora from the east coast of North America (Polychaeta:
Spionidae).—Smithsonian Contributions to Zoology 75:1-32.

574 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1975. Phylum Annelida: Class Polychaeta.—Pp. 153-243 in R. I. Smith and J. T. Carlton,
eds. Light’s Manual: Intertidal Invertebrates of the Central California Coast. 3rd Edition.
University of California Press, Berkeley, 716 pp.

, and J. D. Kudenov. 1978. The Spionidae (Polychaeta) from southeastern Australia and

adjacent areas with a revision of the genera—Memoirs of the National Museum of Victoria

39: 171-280.

Foster, N. M. 1971. Spionidae (Polychaeta) of the Gulf of Mexico and the Caribbean Sea.—Studies
on the Fauna of Curagao and other Caribbean Islands 36: 1-183.

Kudenov, J.D. 1982. The first record of Boccardiella ligerica (Polychaeta: Spionidae) from Imperial
County, California.—Bulletin of the Southern California Academy of Sciences. [submitted
manuscript]

Light, W. J. 1978. Spionidae Polychaeta Annelida.—Invertebrates of the San Francisco Bay Estuary
System. The Boxwood Press, Pacific Grove, California, 211 pp.

Michaelis, H. 1978. Zur Morphologie und Okologie von Polydora ciliata und P. ligni (Polychaeta,
Spionidae).—Helgolander Wissenschaftliche Meeresuntersuchungen 31:102—116.

Rice, S. A., and J. L. Simon. 1980. Intraspecific variation in the pollution indicator polychaete
Polydora ligni (Spionidae).—Ophelia 19:79-115.

Soderstrom, S. 1920. Studien uber die Polychaetenfamilie Spionidae.—Inaugural-Dissertation, Upp-
sala, 286 pp.

Webster, H. E. 1879. The Annelida Chaetopoda of New Jersey.—Annual Report of the New York

Museum of Natural History 32:101-128.

Department of Biological Sciences, University of Alaska, Anchorage, 3221
Providence Drive, Anchorage, Alaska 99508.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 575-578

ASTREPTOS YLLIS ACRASSISETA, A NEW GENUS AND
SPECIES OF THE SUBFAMILY EUSYLLINAE
(POLYCHAETA: SYLLIDAE)

FROM AUSTRALIA

Jerry D. Kudenov and John H. Dorsey

Abstract.—Astreptosyllis acrassiseta is a new genus and species of syllid poly-
chaete belonging to the subfamily Eusyllinae from Victoria, Australia. Astrep-
tosyllis exhibits characters intermediate between those of Streptosyllis and Syl-
lides.

Attempts to confirm the identity of a syllid polychaete originally identified as
Streptosyllis led to a revision of all known Streptosyllis species (Kudenov and
Dorsey, in preparation), and to the present paper describing Astreptosyllis acras-
siseta. The new genus is compared to both Streptosyllis Webster and Benedict,
and Syllides Orsted with which it exhibits intermediate characteristics.

Specimens were collected as part of the Port Phillip Bay and Westernport Bay
Environmental Surveys by the Marine Pollution Studies Group (now known as
the Marine Science Laboratories), Ministry for Conservation, Melbourne, Vic-
toria. Details of the survey areas, stations and their physical parameters are
presented by Poore et al. (1975) and Coleman et al. (1978).

Specimens are deposited at the following institutions: Allan Hancock Foun-
dation, University of Southern California, Los Angeles (AHF); Australian Mu-
seum, Sydney (AM); National Museum of Victoria, Melbourne (NMV); and the
National Museum of Natural History, Smithsonian Institution, Washington, D.C.
(USNM).

Astreptosyllis, new genus

Type-species.—Astreptosyllis acrassiseta new species, by monotypy.

Diagnosis.—Prostomium with 2 pairs of eyes, 3 antennae; paired palps fused
basally, distally digitiform, directed anteriorly and ventrally (visible from above).
Pharynx unarmed with distal circlet of papillae. Proventriculus occupying up to
8 segments. Peristomium with 2 pairs of tentacular cirri, latter long, cylindrical,
pseudoannulate, slightly club-shaped. Dorsal cirri long, cylindrical, pseudoan-
nulate anteriorly, becoming strongly annulate posteriorly. Ventral cirri smooth,
short anteriorly, becoming prolonged posteriorly. Parapodia uniramous, sup-
ported by single aciculum. Acicula slightly knob-shaped distally, having thin-
shafts throughout. Superior dorsal simple setae with distal hoods. Composite
falcigers of 2 kinds, with or without distal hoods: anterior ones having thick-
shafts and short, enlarged blades; and medial and posterior ones having thin-
shafts and narrow blades. Inferior ventral simple setae absent. Pygidium lacking
anal cirri.

Remarks .—Astreptosyllis is most closely allied to Streptosyllis and Syllides. It
is most similar to Streptosyllis in having 2 kinds of composite falcigers, superior

576 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

dorsal simple setae bearing distal hoods, and prolonged ventral cirri in posterior
setigers. Astreptosyllis is also similar to Syllides in having only thin-shafted acic-
ula in all setigers. In this feature, Astreptosyllis clearly differs from Streptosyllis
since the latter possesses two kinds of acicula. Syllides further differs from
Astreptosyllis in having only one kind of composite falciger, in lacking distal
hoods on dorsal simple setae, and in lacking prolonged ventral cirri in posterior
setigers. Only the type-species, described below, is known.

Etymology.—The generic name derives from the Latin prefix ‘a’ meaning with-
out, and the generic name, Streptosyllis. Gender: feminine.

Astreptosyllis acrassiseta, new species
Fig. |

Material examined.—VICTORIA (AUSTRALIA): Westernport, Westernport
Bay Environmental Study Station 1722, 9 m, coarse sand, 22 November 1973;
holotype (NMV G2771), | paratype (NMV G2772). Sta. 1707, 1 m, coarse sand,
7 January 1974; 1 paratype (NMV G2773). Sta. 1724, 18 m, medium sand, 22
November 1973; 1 paratype (NMV G2774). Sta. 1728, 10 m, coarse sand, 20
November 1973; .1 paratype (NMV G2775). Port Phillip Bay, Port Phillip Bay
Environmental Study Station 945/3, 4m, coarse to medium sand, November 1971;
1 paratype (NMV G2776). Sta. 945/4; 1 paratype (AM W.18586). Sta. 973/2, 13
m, medium sand, February 1971; 1 paratype (AM W.18587). Sta. 973/4; 1 paratype
(USNM 74488). Sta. 984/44, 5 m, coarse sand, February 1971; 1 paratype (AHF
POLY 1224). Sta. 985/3, 9 m, coarse sand, December 1971; 2 paratypes (AHF
EOL Y 1223):

Description.—Small species, measuring up to 3.1 mm long, 0.3 mm wide with-
out parapodia, 0.5 mm wide with parapodia, for up to 42 setigers; holotype 2.0
mm long. Body lacking pigmentation in alcohol; articles of annulated dorsal cirri
with golden-yellow pigment granules; with lateral ciliary bands on prostomium,
segments and pygidium.

Prostomium wider than long, with 2 pairs of red-colored eyes in trapezoidal
arrangement, the anterior pair farthest apart; antennae slightly club-shaped and
pseudoannulate, these missing in all specimens except holotype. Lateral antennae
inserted between anterior pair of eyes; median antenna inserted between posterior
pair of eyes as indicated by scars. Palps large, directed anteriorly, becoming
ventrally oriented towards the digitiform tips (Figs. la, b). Pharynx unarmed,
with 10 terminal papillae. Proventriculus large, pear-shaped, generally present in
setigers S—12 [or setigers 3-9 of paratype (NMV G2772)]. Peristomium with 2
pairs of long, cylindrical tentacular cirri, pseudoannulate and slightly club-shaped
distally (Fig. la).

Anterior parapodia truncate distally (Fig. 1c), becoming elongate and conical
posteriorly (Fig. 1d). Dorsal cirri of first 2 setigers resembling antennae and ten-
tacular cirri; those following strongly annulate with anterior ones each having 6
oblong articles, increasing to 9 posteriorly. Ventral cirri of anterior segments
thick, conical, extending to tips of parapodial lobes (Fig. 1c), becoming greatly
prolonged and pseudoannulate posteriorly (Fig. 1d).

Acicula thin-shafted with distally knobbed tips (Fig. le), being difficult to detect
in dense setal fascicles of setigers 1-6. Superior dorsal simple setae present in all

VOLUME 95, NUMBER 3 S17)

N
N
x
N
N
N
\
N

0.05 mm

Fig. 1. Astreptosyllis acrassiseta: a, Holotype: Anterior end, dorsal view, pharynx extended; b,
Paratype (NMV G2772): Anterior end, lateral view; c-k, Holotype: c, Parapodium from setiger 3,
anterior view; d, Parapodium from setiger 29, anterior view; e, Aciculum from setiger 3; f, Dorsal
simple seta from setiger 3; g, Anterior enlarged composite falciger; h, Same, ventral view; i, Posterior
composite falciger; j, Same, detail of blade tip; k, Same, detail of shaft tip, ventral view.

578 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

setigers, having thin-shafts, tapered blunt tips, digitiform to vesiculate hoods (Fig.
If).

Setigers 1-6 each with 8-12 enlarged composite falcigers; blades stout, distally
unidentate with large, rounded basal spurs and smooth cutting margins (Fig. 1g),
appearing distally bifid in ventral view (Fig. lh); shaft tips with dorsal superior
branch terminating in 1 medial tongue-shaped lobe and a broad subterminal cup-
shaped flange, with ventral inferior branch broadly incised distally (Figs. 1g, h).
From setiger 7 to end of body, each fascicle with 6-8 slender composite falcigers;
blades long, unidentate, with funnel-shaped distal hoods and minutely serrated
cutting margins (Figs. li, j); blades of superior falcigers about 2x longer than
lower ones within a fascicle; shaft tips not inflated, resembling those of setigers
1-6 (Figs. li, k). Transition between setigers 6—7 abrupt.

Pygidium wider than long, 2 longer than preanal segments, lacking anal cirri.

Etymology.—The epithet, acrassiseta, refers to the absence of enlarged acic-
ular spines; it is regarded as a noun in apposition.

Acknowledgments

We are grateful to the Marine Science Laboratories (formerly Marine Studies
Group), Ministry for Conservation, Victoria, Australia, for their support of this
research. We are also indebted to the following individuals who have commented
on various drafts of this paper: K. Fauchald and M. Pettibone, National Museum
of Natural History; J. A. Blake, Suffolk University; and W. J. Light, University
of Minnesota. S. J. Williams generously made the resources of the Allan Hancock
Foundation readily available.

Literature Cited

Coleman, N., W. Cuff, M. Drummond, and J. D. Kudenov. 1978. A quantitative survey of the
macrobenthos of Western Port, Victoria.——Australian Journal of Marine and Freshwater Re-
search 29:445-466.

Kudenov, J. D., and J. H. Dorsey. 1982. A revision of the genus Streptosyllis (Polychaeta: Syllidae:
Eusyllinae).—Proceedings of the Biological Society of Washington. [In preparation.]

Orsted, A. S. 1845. Fortegnelse over Dyr, samlede i Christianiafjord ved Drgbak fra 21-24 July
1844.—Naturhistorisk Tidsskrift Kjgbenhavn Series 2, 1:400—427.

Poore, G. C. B., S. F. Rainer, R. B. Spies, and E. Ward. 1975. The zoobenthos program in Port
Phillip Bay, 1969-73.—Victoria Fisheries and Wildlife Paper Number 7, 78 pp.

Webster, H. E., and J. E. Benedict. 1884. The Annelida Chaetopoda from Provincetown and
Wellfleet, Mass.—Annual Report of the United States Commissioner of Fish and Fisheries for
1881:699-747.

(JDK) Department of Biological Sciences, University of Alaska, Anchorage,
3221 Providence Drive, Anchorage, Alaska 99508; (JHD) Interstate Electronics
Corporation, P.O. Box 3117, Anaheim, California 92803.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 579-582

PINNIXA COSTARICANA, A NEW SPECIES
OF CRAB FROM CENTRAL AMERICA
(BRACHYURA: PINNOTHERIDAE)

Mary K. Wicksten

Abstract.—Pinnixa costaricana, a new species of pinnotherid crab, has been
found in Guanacaste Province, Costa Rica. The crab, related to P. floridana and
P. bahamondei, has a very small palpus on the third maxilliped, teeth on the
merus of the third walking leg, and a nearly smooth carapace, without marked
ridges.

The intertidal fauna of western Costa Rica is poorly known. During a recent
collecting trip, Richard C. Brusca brought specimens of decapod crustaceans and
other invertebrates to the Allan Hancock Foundation (AHF), University of
Southern California. Among the specimens were two small crabs belonging to an
undescribed species.

Pinnixa costaricana, new species
Figs. 1, 2

Diagnosis.—Small commensal crab. Carapace 2.3 as wide as long in adult,
anterolateral margin without crest, cardiac ridge lacking. Merus of third walking
leg 1.25x long as wide, dactyls of walking legs 1-3 nearly straight, dactyl of
fourth walking leg short and triangular. Chelae weak, with very tiny granules.
Palpus of third maxilliped small, merus rounded.

Description.—Carapace suboblong, cylindrical, anterolateral margins diverging
posteriorly, forming shoulders from which side walls drop away vertically, sur-
face punctate, slight gastrocardiac trench, no cardiac ridge or anterolateral crest.
Posterior margin straight. Front not advanced, truncate, lobes separated by shal-
low median sulcus. Margins with some setae, especially on ventral surface. Orbits
small, slightly inclined downward.

Third maxilliped with merus having rounded margins, palpus small, dactyl bare-
ly reaching merus and inserted at distal end of propodus, entire appendage setose.

Chelipeds slender, setose. Margins of chelae subparallel, manus with pile, tiny
granules on propodus. Fingers slender, tips pointed and curved, meeting without
gape.

First two walking legs slender, meri trigonal, dactyls sharp, slender, and nearly
Straight. First walking leg somewhat smaller than second, nearly reaching end of
propodus of second leg. Second walking leg reaching carpus of third walking leg.
Third leg very wide; merus 1.25x as long as wide, with | stout tooth and group
of smaller teeth and tubercles at posterodistal angle; carpus without teeth, pro-
podus 1.5x as long as wide, with granules along flexor margin; dactyl straight
and slender. Fourth walking leg short, reaching almost to end of merus of third
leg, dactyl stout and triangular. All legs pilose.

580

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Pinnixa costaricana, holotype. Playa de Coco, Guanacaste Province, Costa Rica.

Fig. 1.

581

VOLUME 95, NUMBER 3

“MOIA [BJUOIJ Ul UOTSAL [eIIGIO ‘q ‘epayD ‘od -‘uswopge sjeulo,y ‘gq ‘podiyixew pry ‘y :edAjojoy ‘vunoupisoo pxmuig

7 SI

582 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Female abdomen widest at third segment, tapering slightly at fourth segment,
fringed with long setae.

Holotype.—Female, carapace length 1.8 mm, width 4.2 mm. Playa de Coco,
Guanacaste Province, Costa Rica (about 10°5’N, 85°45’W), low intertidal zone,
sand and rocks, 27 April 1980, R. C. Brusca, AHF type number 806. Paratype,
juvenile, same location and date, carapace length 1.2 mm, width 2.3 mm, U.S.
National Museum of Natural History.

Discussion.—Pinnixa costaricana resembles P. floridana Rathbun, 1918, P.
pembertoni Glassell, 1935, and P. bahamondei Garth, 1957 in having granules or
teeth on the third walking leg. It differs from the other species in having a very
tiny palpus and a rounded merus of the third maxilliped, instead of a subrectan-
gular merus and long palpus. In P. floridana, from the Gulf of Mexico, the merus
of the third walking leg is 1.66 as long as wide, and the carapace is 2 as long
as wide. Pinnixa pembertoni, from the Gulf of California, has a tooth on the
dactyl of the chela, ridges on the chela, and a gape between the fingers of the
chela. The carapace has a sharp marginal angle at the junction of the anterior and
posterior margins. Pinnixa bahamondei, from Chile, has a tooth on the carpus
of the third walking leg, and more pronounced granules on the chela. The merus
of the third walking leg is 1.5x< as long as wide, and the propodus is as long as
wide. The dactyl of the fourth walking leg is slender, not stout.

The host of P. costaricana was not recorded. The small size and slender shape
of the crab, however, seem adapted for life in the tube or burrow of a polychaete.

Acknowledgments

I thank John S. Garth, University of Southern California, for reviewing the
manuscript. Paula Walker, Texas A&M University, prepared the figures.

Literature Cited

Garth, J. S. 1957. The Crustacea Decapod Brachyura of Chile. Reports of the Lund University
Chile Expedition. 1948-49, Number 29.—Lunds Universitats Arsskrift. N.F. Avd. 2, Bd. 53,
Ihe Yo WAS) jayos

Glassell, S. A. 1935. New or little known crabs from the Pacific coast of northern Mexico.—
Transactions of the San Diego Society of Natural History 8(14):91-106.

Rathbun, M. J. 1918. The grapsoid crabs of America.—Bulletin of the United States National Mu-
seum 97:1-461.

Department of Biology, Texas A&M University, College Station, Texas 77843-
8250"

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 583-593

REVIEW OF SOME LITTLE-KNOWN SPECIES OF SYLLIDS
(ANNELIDA: POLYCHAETA) DESCRIBED FROM THE
GULF OF MEXICO AND CARIBBEAN BY
HERMANN AUGENER IN 1924

Joan M. Uebelacker

Abstract.—The types of 6 little-known syllid species described by Augener in
1924 from the Dry Tortugas, Florida, and from St. Thomas and St. Croix in the
West Indies, were reexamined. Haplosyllides floridana is a sexual form herein
assigned to Haplosyllis floridana n. comb.; Haplosyllis aberrans (Fauvel) is
synonymized with it. Eusyllis antillensis and Syllis (Typosyllis) tigrinoides are
synonyms; the latter name is retained. Syllis (Typosyllis) tortugaensis is referred
to S. (7.) armillaris (Muller). Syllis (Typosyllis) fuscosuturata has previously
been synonymized with Branchiosyllis exilis (Gravier); that synonymy is sup-
ported here. Syllis (Typosyllis) corallicoloides remains a valid species.

' In 1924, H. Augener erected six new species of syllids, but provided only short,
vague descriptions without illustrations. Four of these species were mentioned
in subsequent works by the same author, but again no figures were included.
Later authors, who reported some of these species, did little to augment the
original descriptions, although Monro (1933a, b) provided several figures for Syllis
fuscosuturata and S. tigrinoides, and Westheide (1974) furnished detailed illus-
trations of Branchiosyllis exilis, to which he referred Augener’s S. fuscosuturata.
The remaining four species have never been illustrated.

In preparation for a comprehensive study of Gulf of Mexico syllids, I reex-
amined Augener’s type specimens to compare them with available syllids from
the Gulf. The need for complete descriptions and illustrations of Augener’s species,
along with a discussion of their taxonomic status, became evident and resulted
in this paper.

Branchiosyllis Ehlers, 1887
Branchiosyllis exilis (Gravier, 1900)

Syllis (Typosyllis) exilis Gravier, 1900:160, pl. 9, fig. 9, text-fig. 28-30.

Syllis (Typosyllis) fuscosuturata Augener, 1924:43; 1927b:52.—Hartman, 1959:
230.—Perkins and Savage, 1975:31.

Syllis fuscosuturata.—Monro, 1933a:32, fig. 14; 1933b:250, fig. 4a—c.—Hartman,
1939:10.

Syllis exilis.—Monro, 1937:82, 1939:386.

Branchiosyllis exilis.—Westheide, 1974:60, fig. 26 (synonymy).

Typosyllis fuscosuturata.—Fauchald, 1977:21.

Material examined.—FLORIDA: Tortugas, S.W. Channel, 12 Feb. 1907, 2
syntypes of Syllis fuscosuturata (ZMHUB 6598); Station 321, zone 5, traverse
10, J. S. Colman and G. Tandy, colls., 26 July 1931, C. C. A. Monro, ID, as
Syllis fuscosuturata (BMNH ZK.1932.12.22.137, 1 specimen).

584 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Remarks.—Monro (1937:82) first considered Syllis fuscosuturata a synonym of
S. exilis. Later, Westheide (1974:60) transferred the latter to the genus Bran-
chiosyllis, and included S$. fuscosuturata in synonymy on the basis of material
from Curacao and the West Indies, identified by Augener and deposited in the
Zoologisches Museum der Universitat Hamburg. The syntypes and Monro’s
specimen of S. fuscosuturata examined here agree well in all characteristics with
the descriptions of B. exilis published by Westheide (1974) and Gravier (1900).

Distribution.—Circumtropical.

Haplosyllis Langerhans, 1879
Haplosyllis floridana (Augener, 1924), new combination
Fig. |

Haplosyllides floridana Augener, 1924:44.—Hartman, 1959:208.

Syllis (Haplosyllis) aberrans Fauvel, 1939:290, fig. 3.

Haplosyllis aberrans.—Hartman, 1954:622, 626, 629; 1959:208.—Hartmann-
Schroder, 1978:49, figs. 1-7.—Rullier and Amoureux, 1979:160.

Material examined.—FLORIDA: Tortugas, Hartmeyer, coll., 8 July 1907, ho-
lotype of Haplosyllides floridana (ZMHUB 6608).

Description.—Complete holotype yellowish-brown in color, with 15 setigers,
length 1.8 mm, width at midbody 0.4 mm including parapodia. Prostomium fused
with first segment, anteriorly rounded, with 3 short digitiform antennae (Fig. la).
Eyes, palps, and tentacular cirri lacking. First segment setigerous. Parapodia
biramous, with globular notopodia and slender, prolonged neuropodia. Dorsal
cirri filiform, about 4 times length of parapodia on setiger 1, thereafter alternating
about twice and 1.5 times length of parapodia. Ventral cirri clavate, inserted
midway on ventral surfaces of parapodia and extending about halfway between
tips of notopodia and neuropodia. Notopodia with solitary, slender acicula slightly
bent at tips, and numerous long, capillary natatory setae starting on setiger 2.
Neuropodia with solitary, stout acicula narrowing at tips (Fig. 1b), and 2-3 long,
stout setae with somewhat constricted trifid tips (Fig. 1c). Dorsalmost neurosetae
slightly larger than others. Pygidium rounded, narrower than posterior segments,
lacking cirri. Pharynx and proventricle absent.

Remarks.—The holotype is clearly an epitoke based on the following evidence:
reduction of prostomial appendages, immense pair of eyes (reported by Augener,
now completely faded), absence of peristomium and anterior digestive tract, and
presence of notopodia and natatory setae. According to Augener, the holotype
was possibly collected in the plankton.

Characteristics of the setae and dorsal cirri of Haplosyllis floridana agree well
with H. aberrans (Fauvel). The latter was originally described from Indochina,
associated with sponges and H. spongicola, by Fauvel (1939). Haplosyllis aber-
rans was later reported by Hartman (1954) from the northern Marshall Islands,
associated with coral. Hartmann-Schroder (1978) reported it with ophiuroids from
Puerto Rico, and suggested that its occurrence may be circumtropical. Rullier
and Amoureux (1979) recorded H. aberrans from Brazil on substrates of coral
and calcareous algae. Epitokes have not previously been noted.

Distribution.—Indochina; Enewetak Atoll, Marshall Islands; Puerto Rico; Dry
Tortugas, Florida; Brazil. .

VOLUME 95, NUMBER 3 585

E
E
ie
‘o)

0.02 mm

4 See 5 rs
Ye: st (

‘Fig. 1. Haplosyllis floridana, epitoke: a, Anterior end, dorsal view; b, Aciculum; c, Neuroseta.
From holotype of Haplosyllides floridana (ZMHUB 6608).

Syllis Savigny, 1818
Subgenus 7yposyllis Langerhans, 1879
Syllis (Typosyllis) armillaris
(Muller, 1771, in Muller, 1776)
Fig. 2

Syllis (Typosyllis) tortugaensis Augener, 1924:43; 1927b:51.—Hartman, 1959:
231.—Perkins and Savage, 1975:31.
Typosyllis tortugaensis.—Hartman, 1959:235.

Material examined.—FLORIDA: Tortugas, Bird Key, syntype of S. tortu-
gaensis (ZMHUB 6596).

Description.—Complete syntype grayish-brown in color, with 198 setigers, length
30 mm, width in proventricular region 0.9 mm including parapodia. Body arched
dorsally, flattened ventrally. Prostomium anteriorly rounded, posteriorly nearly
linear (Fig. 2a). Eyes 4, faded, anterior pair larger, lentigerous. Median antenna
inserted between anterior eyes, with about 17 articles. Lateral antennae inserted
near anterior margin of prostomium, two-thirds length of median antenna, with
13-18 articles. Palps bluntly rounded anteriorly, apparently not fused basally,
shorter than lateral antennae. Nuchal organs not observed. Peristomium slightly
shorter than following segments. Left dorsal tentacular cirrus missing, right one
subequal in length to median antenna, with about 17 articles. Ventral tentacular
cirri subequal in length to lateral antennae, with about 14 articles. Segments about
4 times broader than long anteriorly. Parapodia distally rounded (Fig. 2b), oc-
casionally with small pre- and postsetal lobes. Dorsal cirri fairly stout and dis-
tinctly articled throughout, cirriform anteriorly, becoming fusiform in mid-body
region (Fig. 2b), shorter than body width except on anterior setigers, alternating

586 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

danse f g h

Riga 2aySyliis Clk yposyllis) armillaris: a, Anterior end, dorsal view; b, Parapodium from mid-body
region, anterior view; c, Acicula from anterior region; d, Superior falciger from anterior region; e,
Inferior falciger from anterior region; f, Superior falciger from mid-body region; g, h, Inferior falcigers
from mid-body region; i, j, Inferior falcigers from posterior region; k, Superior simple seta; 1, Inferior
simple seta. Scale same for d—l. From syntype of Syllis tortugaensis (ZMHUB 6596).

VOLUME 95, NUMBER 3 587

slightly in length, longer ones with about 15-30 articles, shorter ones with about
10-20 articles. Ventral cirri digitiform, generally not extending beyond tips of
parapodia. Acicula sometimes slightly emergent, numbering 5 in anterior para-
podia (Fig. 2c), 3 in middle parapodia, 2 stout and | very slender (Fig. 2b). Setae
mostly composite falcigers with serrate shaft-heads. Setae of anterior region bi-
dentate, with numerous short serrations below subdistal tooth, blades graded in
length from superior (Fig. 2d) to inferior (Fig. 2e), blade length ratio* 1.5:1. Setae
of mid-body region including superior bidentate falcigers with small subdistal
tooth (Fig. 2f), and inferior subbidentate to unidentate falcigers (Fig. 2g, h), blade
length ratio 1.2:1. Setae of posterior region again bidentate, some similar to an-
terior setae (Fig. 21), some more slender (Fig. 2j), blade length ratio 1.4:1; each
posterior parapodium with single, short, slender, minutely serrate superior and
inferior simple setae (Fig. 2k, 1). Paired anal cirri missing; short, slender, smooth,
midventral anal cirrus present. Pharynx occupying setigers 1-11, middorsal tooth
subterminal. Proventricle occupying setigers 12—24, with 44 rows of muscle cells,
4 times longer than wide and 1.2 times longer than pharynx.

Remarks.—Syllis tortugaensis is herein referred to S. armillaris, based on the
presence of fusiform dorsal cirri, and falcigers which are bidentate anteriorly and
posteriorly, and unidentate or subbidentate in the mid-body region. A specimen
(BMNH ZK.1932.12.22.124) from the Dry Tortugas identified by Monro (1933b:
251, fig. 5) as S. tortugaensis does not belong to S. armillaris, but may represent
an undescribed species.

Distribution.—Cosmopolitan.

Syllis (Typosyllis) corallicoloides Augener, 1924
Fig. 3

Syllis (Typosyllis) corallicoloides Augener, 1924:42; 1927a:134; 1927b:51; 1933:

227; 1936:341.—Hartman, 1959:230.—Perkins and Savage, 1975:31.
Typosyllis corallicoides (sic).—Hartman, 1951:41.—Marsden, 1960:995.
Typosyllis corallicoloides.—Hartman, 1959:234.

Material examined.—WEST INDIES: St. Croix, Grube, coll., 2 syntypes
(ZMHUB 3020).

Description.—Body yellowish-brown in color, nearly complete larger syntype
with 110 setigers, incomplete smaller syntype somewhat desiccated, with 76 se-
tigers, lengths 17.7 and 15.2 mm, respectively, widths in proventricular region
1.25 and 0.55 mm including parapodia. Body flattened ventrally, dorsally arched
anteriorly becoming more flattened in mid-body region. Prostomium suboval to
rounded. Eyes 4, small, very faded, in trapezoidal arrangement with anterior pair
farther apart (Fig. 3a), not visible on larger syntype. Antennae, tentacular and
dorsal cirri much longer than body width, with flattened, twisted, indistinct ar-
ticles appearing smooth under low magnification on larger syntype, but more
distinct on smaller syntype; all article counts given for smaller syntype only.
Median antenna missing, ceratophore inserted between eyes. Lateral antennae
with about 15 articles. Palps elongate, blunt anteriorly, about half length of lateral

* Ratio of lengths of longest superior to shortest inferior setal blades.

588 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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Fig. 3. Syllis (Typosyllis) corallicoloides, smaller syntype (ZMHUB 3020): a, Anterior end, dorsal
view; b, Parapodium from anterior region, posterior view; c, Acicula from anterior region; d, Acicula
from mid-body region; e, Superior falciger from anterior region; f, Inferior falciger from anterior
region; g, Superior falciger from mid-body region; h, Inferior falciger from mid-body region; i, Su-
perior simple seta; j, Inferior simple seta. Scale same for e-j.

VOLUME 95, NUMBER 3 589

antennae. Nuchal organs small, paired lobes between prostomium and peristo-
mium. Peristomium shorter than following segments, forming small flap over
posterior part of prostomium. Dorsal tentacular cirri with 24 articles, ventral with
about 17 articles. Parapodia with short pre- and postsetal lobes on larger syntype,
distally rounded on smaller syntype (Fig. 3b). Dorsal cirri elongate, cirriform
anteriorly, becoming more filiform in middle and posterior regions, alternating
somewhat in length, longer cirri with 30-35 articles, shorter cirri with 18—24
articles. Ventral cirri digitiform, extending beyond tips of parapodia anteriorly.
Acicula with tips emergent, numbering 4 in anterior parapodia (Fig. 3c), 2 in mid-
body region (Fig. 3d). Setae mostly composite, bidentate falcigers (Fig. 3e—h),
with lightly serrate shaft-heads. Blades graded in length from superior to inferior,
teeth rounded or pointed, with fine, short serrations below subdistal tooth. Setae
fairly slender anteriorly (Fig. 3e, f), becoming somewhat broader in mid-body
region (Fig. 3g, h). Posterior falcigers essentially similar to those of mid-body
region, but with slightly shorter blades and with 1-2 more slender superior fal-
cigers in posteriormost parapodia. Setal blade length ratios 1.5—1.9:1 in anterior
body region, 1.5—1.6:1 in middle and posterior regions. Posterior parapodia with
additional single, slender, minutely bifid superior and inferior simple setae (Fig.
31, j). Pygidium with paired, posterolateral anal cirri having about 27 articles,
midventral cirrus not observed. Pharynx occupying first 14 setigers in larger syn-
type and setigers 3-10 in smaller syntype, with smooth margin surrounded by
14-16 soft papillae, and with fairly large, subterminally located middorsal tooth.
Proventricle disintegrated in larger syntype, occupying setigers 11-17 in smaller
syntype, with about 35 rows of muscle cells, 3.1 times longer than wide and 1.1
times longer than pharynx.

Remarks.—Syllis corallicoloides differs from most other members of the genus,
with the exception of S. tigrinoides, in having the margin of the pharynx sur-
rounded by more than ten soft papillae. Syllis corallicoloides differs from S.
tigrinoides in having more slender dorsal cirri anteriorly; setal shafts more slender
in the mid-body region, and teeth of the setal blades more separated and less
rounded; greater disparity among lengths of setal blades; fewer papillae surround-
ing the margin of the pharynx; and the proventricle about 3 rather than 4—5 times
longer than wide. Overall similarity between the types of the two species is great;
collection and study of additional specimens may indicate that they represent the
Same species.

Distribution.—West Indies, including St. Croix, Jamaica, Curacao and Bar-
bados; Veracruz, Mexico; Dry Tortugas, Florida.

Syllis (Typosyllis) tigrinoides Augener, 1924
Fig. 4

Syllis (Typosyllis) tigrinoides Augener, 1924:43; 1927b:52; 1931:286; 1933:229.—
Hartman, 1959:231.—Perkins and Savage, 1975:31 (not Monro, 1933b:247, fig.
2a—C).

Typosyllis tigrinoides.—Hartman, 1959:235.,

Eusyllis antillensis Augener, 1924:44.—_Hartman, 1959:204.—Perkins and Savage,
1975:30.

Material examined.—FLORIDA: Dry Tortugas, Bird Key Reef and S.W.
Channel, Hartmeyer, coll., holotype of Syllis tigrinoides (ZMHUB 6597).—WEST

590 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

) Y)
YY)
Up,
Y

f

Fig. 4. Syllis (Typosyllis) tigrinoides: a, Anterior end, dorsal view (dorsal cirri not shown); b,
Parapodium from anterior region, anterior view; c, Parapodium from mid-body region, anterodorsal
view; d, Acicula from anterior region; e, Acicula from mid-body region; f, Superior falciger from
anterior region; g, Inferior falciger from anterior region; h, Superior falciger from mid-body region;
i, Inferior falciger from mid-body region; scale same for d, e, h, i. a, d, e, h, i from holotype of
Eusyllis antillensis (ZMHUB 6594); b, c, f, g from holotype of Syllis tigrinoides (ZMHUB 6597).

VOLUME 95, NUMBER 3 SwIll

INDIES: St. Thomas Sound, Ktikenthal and Hartmeyer, colls., Jan. 1907, ho-
lotype of Eusyllis antillensis (ZMHUB 6594).

Description.—Holotype of S. tigrinoides with dorsum chocolate brown in col-
or, dorsal cirri tan; holotype of E. antillensis yellowish-brown. Both holotypes
incomplete with 104 and 98 setigers, respectively, lengths 17.1 and 15.5 mm,
widths in proventricular region 1.8 and 1.35 mm including parapodia. Body ven-
trally flattened to concave, dorsally arched anteriorly becoming more flattened
in mid-body region. Prostomium suboval to quadrangular (Fig. 4a). Eyes 4, len-
tigerous, in trapezoidal arrangement across posterior half of prostomium. Median
antenna fairly long, on short ceratophore between eyes, with about 45 small,
crowded articles. Lateral antennae half length of median antenna, with about 25
articles. Palps broad, bluntly rounded anteriorly, not fused basally. Nuchal organs
paired, oval lobes covered by peristomial margin. Peristomium about same length
as following segments. Dorsal tentacular cirri slightly longer than median antenna,
with about 45-50 articles. Ventral tentacular cirri slightly longer than lateral an-
tennae, with about 25-30 articles. Segments crowded, up to 8 times broader than
long. Parapodia well developed, slightly larger in mid-body region than anteriorly,
with short, digitiform pre- and postsetal lobes (Fig. 4b, c). Dorsal cirri long and
more or less distinctly articled throughout, mostly curled in holotype of S. tigri-
_noides, becoming more slender towards posterior region, alternating in length,
longer cirri with about 60-75 articles, shorter cirri with about 30-35 articles.
Ventral cirri clavate to digitiform, arising midway on and extending slightly be-
yond tips of parapodia anteriorly, arising basally and not extending beyond tips
of parapodia towards mid-body region. Acicula emergent, numbering 5 in anterior
parapodia (Fig. 4d), 2 stout and 1 minute in middle parapodia (Fig. 4e). Setae
composite falcigers (Fig. 4f-1) numbering about 8-17 in anterior parapodia, about
6—11 in middle parapodia. Shaft-heads with numerous fine serrations in 2—4 trans-
verse rows. Blades graded in length from superior to inferior, bidentate, teeth
usually well-rounded, occasionally pointed, with fine serrations below subdistal
tooth. Setae fairly slender anteriorly (Fig. 4f, g), inferior setae becoming much
stouter in mid-body region (Fig. 41). Setal blade length ratio 1.3—1.5:1 in anterior
region, |.2—1.3:1 in mid-body region. Superior and inferior simple setae not ob-
served (both holotypes incomplete). Pharynx extending back into setigers 13-15,
partially everted in holotype of E. antillensis, somewhat convoluted in holotype
of S. tigrinoides, margin smooth, surrounded by 19-20 soft papillae, middorsal
tooth subterminal. Proventricle rectangular, extending back to setigers 29-30,
with 36—38 rows of muscle cells, 4—S times longer than wide and 1.1—1.5 times
longer than pharynx.

Remarks .—Eusyllis antillensis is herein transferred to the genus Syllis, since
the antennae, tentacular and dorsal cirri are distinctly articled, the palps are not
fused basally, and the margin of the pharynx is smooth rather than denticulate
as in Eusyllis. The holotype of E. antillensis is nearly identical in all respects to
that of S. tigrinoides; the two are hereby synonymized, with the name ftigrinoides
to be retained, since it has page priority in the original publication and has been
more frequently reported in subsequent literature. Three specimens (BMNH
ZK.1932.12.22.180-181) from the Dry Tortugas identified by Monro (1933b:247,
fig. 2a—c) as S. tigrinoides do not belong to this species, but may represent an
undescribed species.

Sy) PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Distribution.—West Indies, including St. Thomas, Curacao, and Jamaica; Dry
Tortugas, Florida; Brazil.

Acknowledgments

I am indebted to Dr. G. Hartwich, Zoologisches Museum an der Humboldt-
Universitat zu Berlin (ZMHUB), for the loan of type material; to A. I. Muir,
British Museum (Natural History) (BMNH), for the loan of specimens; to Ben
F. Jordan, who provided inked renderings of the illustrations; to Thomas H.
Perkins, Florida Department of Natural Resources, for the loan of literature; and
to Dr. Marian H. Pettibone, Smithsonian Institution, for critical review of this

paper.
Literature Cited

Augener, H. 1924. Uber litorale Polychaeten von Westindien.—Sitzungsberichte der Gesellschaft

Naturforschender Freunde zu Berlin, Jahrgang 1922:38—53.

1927a. Polychaeten von Neu-Pommern.—Sitzungsberichte der Gesellschaft Naturforschen-
der Freunde zu Berlin, Jahrgang 1926:119-152.

1927b. Bijdragen tot de Kennis der Fauna van Curagao. Resultaten eener Reis van Dr. C.
J. van der Horst in 1920. Polychaeten von Curagao.—Jaarboekje der Kooniglijk Zoologisch
genootschap ‘‘Natura artis magistra’’ (Amsterdam) 25:39-82.

1931. Die bodensassigen Polychaten nebst einer Hirudinee der Meteor-Fahrt.—Mitteilungen
aus dem Zoologischen Staatsinstitut und Museum Hamburg 44:279-313.

——. 1933. Polychaeten aus den zoologischen Museen von Leiden und Amsterdam. Pt. 1.—

Zoologische Mededeelingen 15:177—260.

1936. Zoologische Ergebnisse einer Reise nach Bonaire, Curacao und Aruba im Jahre 1930.
Polychaeten aus den marinen Salinen von Bonaire und Curagao.—Zoologischen Jahrbucher
(Abteilung fiir Systematik, Okologie und Geographie der Tiere) 67:337-352.

Ehlers, E. 1887. Report on the annelids of the dredging expedition of the U.S. Coast Survey Steamer
Blake.—Memoirs of the Museum of Comparative Zoology at Harvard College 15, vi and 335
pp. + 60 pls.

Fauchald, K. 1977. Polychaetes from intertidal areas in Panama, with a review of previous shallow-
water records.—Smithsonian Contributions to Zoology 221, 81 pp.

Fauvel, P. 1939. Annélides polychetes de I’Indochine recueillies par M. C. Dawydoff.—Commen-
tationes Pontificia Academia Scientiarum, anno 3, 3(10):243-368.

Gravier, C. 1900. Contribution a l’étude des Annélides polychétes de la mer Rouge. Pt. 1.—Nou-
velles Archives du Muséum dhistoire naturelle Paris, Ser. 4, 2(2):137—282.

Hartman, O. 1939. The polychaetous annelids collected on the presidential cruise of 1938.—Smith-

sonian Miscellaneous Collections 98(13):1—22.

1951. The littoral marine annelids of the Gulf of Mexico.—Publications of the Institute of
Marine Science, University of Texas 2(1):7-124.

1954. Marine annelids from the northern Marshall Islands.—United States Geological Sur-
vey, Professional Papers 260 Q:615-644.

1959. Catalogue of the polychaetous annelids of the world. Pts. I and IJ.—Allan Hancock
Foundation Occasional Papers 23, 628 pp.

Hartmann-Schroder, G. 1978. Einige Sylliden-Arten (Polychaeta) von Hawaii und aus dem Kari-
bischen Meer.—Mitteilungen Hamburg Zoologisches Museum und Institut 75:49-61.

Langerhans, P. 1879. Die Wurmfauna von Madeira.—Zeitschrift fiir Wissenschaftliche Zoologie 32:
513-592 + 3 pls.

Marsden, J. R. 1960. Polychaetous annelids from the shallow waters around Barbados and other
islands of the West Indies, with notes on larval forms.—Canadian Journal of Zoology 38:989-
1020.

Monro, C. C. A. 1933a. The Polychaeta Errantia collected by Dr. C. Crossland at Colon, in the
Panama region, and the Galapagos Islands during the expedition of the S. Y. “St. George.’ Pt.
1.—Proceedings of the Zoological Society of London for 1933: 1-96.

VOLUME 95, NUMBER 3 593

1933b. Ona collection of Polychaeta from Dry Tortugas, Florida.—Annals and Magazine

of Natural History, London (10) 12:244—269.

1937. A note on a collection of Polychaeta from the eastern Mediterranean, with the de-
scription of a new species.—Annals and Magazine of Natural History, London (10)19:82-86.
1939. On some tropical Polychaeta in the British Museum mostly collected by Dr. C.

Crossland at Zanzibar, Tahiti and the Marquesas. II. Families Syllidae and Hesionidae.—

Novitates Zoologicae 41:383-393.

Miller, O. F. 1776. Zoologica Danica. Prodromus seu animalium Daniae et Norvegiae indigenarum
characteris, nomine et synonyma imprimis popularium. Copenhagen, xxxiii and 274 pp.
Perkins, T. H., and T. Savage. 1975. A bibliography and checklist of polychaetous annelids of
Florida, the Gulf of Mexico, and the Caribbean region.—Florida Marine Research Publications

No. 14:1-62.

Rullier, F., and L. Amoureux. 1979. Polychaetous Annelids. In Résultats Scientifiques des Cam-
pagnes de la Calypso. Fascicule XI.—Annales de I’Institut Océanographique, N.S. 55, Fas-
cicule Supplémentaire: 145-206.

Savigny, J. C. 1818. Les Annélides. In Lamarck, J. B. de, 1818, Histoire naturelle des animaux
sans vertebres. Paris (Deterville et Verdiere), 5:274-374.

Westheide, W. 1974. Interstitielle Fauna von Galapagos. XI. Pisionidae, Hesionidae, Pilargidae,

Syllidae (Polychaeta).—Mikrofauna Meeresbodens 44: 195-338.

Barry A. Vittor & Associates, Inc., 8100 Cottage Hill Road, Mobile, Alabama
36609.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 594-601

THE STATUS OF TWO CHILEAN FROGS OF THE GENUS
EUPSOPHUS (ANURA: LEPTODACTYLIDAE)

J. R. Formas and M. Inés Vera

Abstract.—External and internal morphological features show that Eupsophus
calcaratus (Gunther, 1881) is a well defined species different from E. roseus (D.
& B.), with which it was previously synonymized. Eupsophus calcaratus 1s re-
described, and notes on its natural history are presented. Eupsophus roseus in-
sularis (Philippi, 1902) is shown to be not conspecific with E. roseus and is
redescribed as Eupsophus insularis.

Gunther (1881) described Cacotus calcaratus, based on only one specimen
from Chiloé Island (no precise locality). Boulenger (1882) transferred this species
to the genus Borborocoetes, and Cei (1958), without an osteological study, con-
cluded that the genera Borborocoetes, Cystignathus, and Alsodes were synony-
mous with the genus Eupsophus. A series of Lynch’s papers (1971, 1972, 1978),
which were based on osteology (especially the cranial bones), permitted the def-
inition of Eupsophus in which two species were included, E. vittatus and E.
roseus; E. migueli Formas, 1978, was not considered by that author. Cei (1958)
and Capurro (1958) considered E. calcaratus a valid species, but they pointed
out that the taxonomic status of this frog needed to be revised. Later, Cei (1960,
1962a, 1962b) and Grandinson (1961) discussed the taxonomy of E. calcaratus
and concluded that the species was synonymous with E. roseus (D. & B.). Formas
(1980) described the karyotype of frogs from Valdivia (Cordillera Pelada) and
Osorno (La Picada) and the name Eupsophus calcaratus was used for these
animals.

During our herpetological studies of Chiloé Island, Osorno, and Valdivia, one
of us (JRF) collected frogs which agree with Guinther’s description and with the
external morphology of the holotype. An osteological study (especially of cranial
osteology) of the Chiloé Island frogs confirms that these animals should be in-
cluded in the genus Eupsophus because they have the cranial pattern of E. roseus,
the type species of the genus (see Lynch 1971). Both E. roseus and E. calcaratus
have the frontoparietal fontanellae large, the sphenethmoid short and extending
between the nasals, which are intermediate in size, separated medially; quadra-
tojugal present; pars facialis of maxillary wide, and columella present.

Eupsophus calcaratus differs from E. roseus in the pigmentation of the upper
part of the iris, the slope of the tips of toes and fingers, distance between pre-
vomerine teeth, and shape of the dorsal and lateral snout profile.

Philippi (1902) described Cystignathus (Borborocoetus) insularis based on five
specimens from Mocha Island, 38°22’S, 73°56’W, 40 km west of the coast of
Arauco Province, Chile. Cei (1958) considered this species synonymous with
Eupsophus roseus, and Capurro (1963), on the basis of only one female from the
same island, concluded that Philippi’s species was a subspecies of Eupsophus
grayi and called the frog Eupsophus grayi insularis. We had the oportunity to

VOLUME 95, NUMBER 3 595

Fig. 1. Adult females of Eupsophus calcaratus from Puntra, Chiloé Island, x<1.5 (left), and E.
insularis from Mocha Island, <1.2 (right).

analyze three specimens (MUZUC 5256, 5299, 5309) from Mocha Island, cour-
teously loaned by Jorge Artigas of the Instituto de Zoologia, Universidad de
Concepcion. We consider that the form described by Philippi is a valid species
because it differs from E. roseus in color pattern, shape of the xiphisternum,
length of fingers and toes, and location of prevomerine teeth. The skull osteology
permits inclusion of Philippi’s taxon into the genus Eupsophus because it shares
the cranial pattern of E. roseus.

Eupsophus calcaratus (Gunther, 1881)
Fig. 1

Cacotus calcaratus Ginther, 1881:19 (Holotype: BMNH 468, 68.9.22.8. Chile;
type-locality: Chiloé Island).

Borborocoetes calcaratus.—Boulenger, 1882:256, pl. 17, Fig. 1.

Cystignathus oxyglossus Philippi, ?1902:110 (Chile; type-locality: eastern part of
Chiloé Island).

Cystignathus fernandezi Philippi, ?1902:112 (Chile; type-locality: Chiloé Island).

Eupsophus calcaratus.—Capurro, 1958:293.—Cei, 1958:270.—Formas, 1980:1163.

Eupsophus grayi Cei, 1960:3; 1962:33.

Eupsophus roseus Grandinson, 1961:128.

Diagnosis.—Eupsophus calcaratus is a moderate sized species that can be
distinguished from its congeners by the following combination of characters: 1)
upper part of the iris bronze-yellow; 2) distal portion of the fingers and toes
rounded and prominent; 3) prevomers in narrow contact; 4) snout pointed in
dorsal and lateral view, noticeably protruding over the lower jaw.

596 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

E. calcaratus E. roseus

Fig. 2. Lateral profile of the snout of Eupsophus calcaratus and E. roseus.

Description (based on 25 living specimens and 31 fixed specimens).—Head
wider than long. Snout pointed in dorsal view, rounded in lateral profile (Fig. 2);
canthus rostralis well defined; loreal region concave, sloping slightly to lip; nos-
trils lateral, closer to tip of snout than to orbit; length of eye greater than distance
between eye and nostril; interorbital distance slightly narrower than length of
eye, greater than internarial distance. Tympanic membrane small and poorly de-
fined, tympanum diameter smaller than eye diameter. Lateral fold present, from
behind eye to middle part of body. Tongue rounded, slightly cordiform. Small
round choanae; dentigerous process of prevomers lying slightly below choanae;
closer medially, transverse or slightly oblique, each process bearing 4—6 teeth.
Forelimbs thin, first finger equal in length to second; third finger much longer
than fourth; digital length in decreasing order 3—4—2—-1. Tips of fingers rounded
and slightly prominent (Fig. 3). Two palmar tubercles ovoid and prominent; sub-
articular tubercles rounded and moderate prominent; supernumerary palmar tu-
bercles absent. Hind limbs slender, tibiotarsal articulation reaching to middle of
eye. Toes long, slender, not fringed; in decreasing order of length 4—(5,3)—2-1.
Outer metatarsal tubercle ovoid and prominent, inner metatarsal tubercle small.

Fig. 3. Lateral profile of the fourth toe of Eupsophus roseus (A) and E. calcaratus (B).

VOLUME 95, NUMBER 3 597

Table 1.—Measurements (mm) of Eupsophus calcaratus from Puntra (Chiloé Island) population.

7 males 24 females
Character Range x +S.D. Range x+S.D.
Snout—vent length 31.4-35.2 32.9 + 1.2 41.2-33.7 Spek as: 20)
Thigh length 15.1-17.0 16.0 + 0.6 16.0—20.2 18.0 + 1.0
Tibia length 16.6—-18.8 17.4 = 0.9 18.1—20.7 19.5 + 0.6
Foot length 23 .2-27.9 24.8 + 1.5 25.0-29.8 28.0 + 0.6
Head width 12.6-14.6 13°52 O28 13.3-15.9 14.5 + 0.6
Head length 11.0-12.2 11.4 + 0.4 11.7-13.8 12.9 + 0.5
Eye diameter 3.2-4.3 3.7 + 0.4 3.64.7 4.1 + 0.3
Eye—nostril distance 2.8-3.1 SAU) ae (sil 3.0-3.8 3.3 + 0.2
Interorbital distance 2.6-3.3 2) z= (3) 2.44.1 3.0 + 0.4
Nostril-snout distance 1.7-2.3 2.0 + 0.2 1.8-2.7 Dep), se (V7
Tympanum diameter 1.8-2.3 2.0 + 0.2 1.9-3.5 2.4 + 0.4

Subarticular tubercles large, conical and prominent; supernumerary tubercles ab-
sent. Tarsal fold absent. Rudiment of web between fourth and third, and between
third and second toes. Anal opening oriented transversely at dorsal level of thighs.
Table 1 shows the variation of external measurements in this species.

Skin smooth, minute tubercles on head. Ventral surface of thighs without gran-
ular tubercles, sometimes with minute tubercles. Two short cutaneous spurs at
the heels and two dorsal linear ridges convergent behind, which are hardly no-
ticeable. Minute tubercles on posterior part of flanks. Dorsum dark gray, with a
remarkable pattern like a butterfly or hourglass extending from head, between
eyes, to middle of dorsum. On flanks a dark fringe reaching the eyes. Some
Specimens show a narrow mid-line. Minute dark spots lateral to the butterfly
pattern; two rounded dark brown spots on lumbar area stand out on the back-
ground. Minute dark spots on whitish venter, most numerous on throat. Arms,
thighs, shanks, and tarsi barred dark brown. In life, dorsum and limbs light brown,
tan or gray, venter whitish. Markings on dorsum, venter and limbs dark brown
or black, flanks sometimes light brown or dark orange and the spots brown. Upper
part of iris bronze-yellow, lower part dark brown.

Eupsophus insularis (Philippi, 1902)
Fig. |

Cystignathus (Borborocoetus) insularis Philippi, 1902:89 (Chile; type-locality:
Mocha Island). (Holotype lost)

Eupsophus grayi Cei, 1958:271.

Eupsophus grayi insularis Capurro, 1963:5—18.

Diagnosis.—Eupsophus insularis is a moderate sized species that can be dis-
tinguished from it congeners by the following combination of characters: 1) dark
brown in color with irregular yellow spots on the dorsum and legs; 2) xiphisternum
truncated and slightly notched; 3) prevomerine teeth below the choanae.

Description (based on 3 fixed specimens).—Head wider than long. Snout slight-
ly truncate in lateral profile and pointed in dorsal view; canthus rostralis well
defined; loreal region flat. Nostrils lateral, closer to tip of snout than to orbit;

598 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Measurements (mm) of Eupsophus insularis.

MUZUC no.

Character 5256 5309 $299
Snout—vent length 40.90 38.20 38.70
Thigh length 20.50 18.10 17.85
Tibia length 22.00 DBs) 20.15
Foot length 32.65 32.65 27.65
Head width 15.45 16.10 15.60
Eye diameter 5225) 4.30 4.25
Eye-—nostril distance 3.20 3.45 3.20
Interorbital distance 3.45 3.70 3.30
Nostril—snout distance 3.00 2.05 DAs)
Tympanum diameter 2.85 i) 2.05

length of eye greater than distance between eye and nostril; internarial distance
slightly smaller than interorbital distance. Tympanum diameter half the eye di-
ameter. Well developed fold extending from behind eye to insertion of arm. Tongue
rounded, without notch at the tip, posterior two thirds free. Small, round choanae;
dentigerous process of prevomers lying medial and below choanae, separated
medially, slightly oblique, each process bearing 6—7 teeth. Forelimbs thin; first
finger equal in length to second; third finger much longerthan fourth; digital length
in decreasing order 3—-4—2—1. Palmar webbing absent; tips of fingers slightly round-
ed and prominent. Inner median palmar tubercle ovoid; outer palmar tubercle
round; subarticular tubercles rounded and of moderate size; supernumerary pal-
mar tubercles absent. Hind limbs slender. Toes long, slender, and moderately
fringed; tips of toes rounded; third and fifth toes equal in length, toes in decreasing
order of length 4—(3,5)—2—1. Outer metatarsal tubercle small and rounded; inner
metatarsal tubercle ovoid and prominent. Subarticular tubercles prominent and
rounded; supernumerary tubercles absent. Tarsal fold absent. Toes without web-
bing. Anal opening oriented transversely, at dorsal level of thighs.

Skin smooth on dorsal and ventral surfaces; posteroventral areas of thighs with
flat tubercles. Dorsum and legs brown with yellow, minute, irregular spots; venter
yellowish and throat with dark brown irregular spots. Color of ventral surface of
thighs similar to the belly. Arms marbled.

Variation.—Table 2 summarizes the variation in proportions of the three spec-
imens examined. Two (MUZUC 5256, 5299) of the three specimens examined
show the same color pattern, but the third (MUZUC 5309) has large, yellow,
irregular spots on the dorsum, arms and shanks; and a light triangle on the head
that extends from the interorbital area to the snout. The belly and the throat of
this specimen are light brown.

Comparisons

In habitus, size, and dorsal pattern, E. calcaratus is similar to E. roseus;
however the species show external differences. Eupsophus calcaratus has the
upper part of the iris bronze-yellow colored and the belly is whitish with minute,
irregular dark spots; E. roseus has the ventral area transparent or whitish and

VOLUME 95, NUMBER 3 599

ul
3
3

Fig. 4. Pectoral girdle of Eupsophus insularis.

the upper part of the iris is orange colored. Eupsophus calcaratus and E. migueli
have the same color in the upper part of the iris, but the species are different in
ventral color pattern; the belly of E. migueli is wine with irregular white spots.
Eupsophus insularis has a dorsal color pattern (dark brown with yellow irregular
spots) that is not present in the other species of the genus; this agrees with the
color description given by Capurro (1963). This species has the xiphisternum
slightly notched as in E. migueli, but in the latter the notch is greater (Fig. 4).
On the basis of this character, E. insularis and E. migueli differ from the re-
maining species of the genus, EF. vittatus and E. roseus. In E. insularis the pre-
vomerine teeth are much more below the choanae than in the other Eupsophus
species. Eupsophus insularis is a frog of moderate size (x = 39.3 mm) as are E.
calcaratus (x = 35.1 mm), E. roseus (x = 36.0 mm; Cei 1962) and E. migueli
(x = 35.5 mm; Formas 1978). All these are remarkably smaller than E. vittatus
(x = 59.4 mm; Grandinson 1961).

Twenty skulls of E. roseus and E. calcaratus and one skull of E. insularis were
examined. Although E. roseus, E. insularis and E. calcaratus share the same
cranial pattern (Fig. 5), qualitative differences were detected. The latter species
has the prevomer bones in median contact and the sphenethmoid extends anterior
to the nasals. The nasal bones show a tendency to be in median contact. The

600 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

10mm

Fig. 5. Dorsal and ventral views of the skulls of Eupsophus roseus (A), E. calcaratus (B) and E.
insularis (C).

prevomers of E. roseus are separated and the sphenethmoid reaches and some-
times slightly exceeds the anterior border of the nasals. The anterior end of the
cultriform process of E. insularis does not reach between the palatines.

Natural History

Observations of E. calcaratus were made at Puntra, Chiloé Island (42°03’S,
73°48'W). A male was observed at night when it was calling at the border of a
stream (air temperature 5°C) on 15 September 1980. In the same place we also
recorded the mating call of E. vittatus. Females (8) with white-creamy ovarian
eggs (59-168) were found on 28 July 1981. Adults males, females, and juveniles
were collected under logs in a Chiloé Nothofagus forest on 15 September 1980
and found under logs outside the forest on 28 July 1981. Other herpetozoa found
in the same area were the small lizard Liolaemus pictus and the frogs E. vittatus,
Batrachyla taeniata and Batrachyla leptopus.

The three specimens of E. insularis are females, collected at the end of summer
(14 March 1971). Two of these frogs have creamy-white ovarian eggs (119 in
MUZUC 5256). MUZUC 5309 is immature.

Specimens Examined

Abbreviations.—Instituto de Zoologia, Universidad Austral de Chile (IZUA),
Museo de Zoologia, Universidad de Concepcion (MUZUC).

Eupsophus calcaratus: Chiloé Island: IZUA 1826-1856, Puntra; 1888-1907
(skeletons), Yaldad; 1908, Caulin; 1909-1910, Huillinco; 1911, Coquiao; 1912—
1914, San Juan; 1916-1917, Quetalco; 1918, Mocopulli; 1919, Castro; 1920, Terao.

VOLUME 95, NUMBER 3 601

Valdivia Province: 1779-1783, Cordillera Pelada. Osorno Province: 973, 1582—
1583, La Picada; 1754-1755, Antillanca.

Eupsophus roseus: San Martin Forest, Valdivia Province; IZUA 1857-1887;
1529-1533 (skeletons); 1691-1692 (skeletons); 1964—1968 (skeletons); 1700-1702
(skeletons); 1922—1926 (skeletons).

Eupsophus insularis: Mocha Island: MUZUC 5256, 5309, 5299.

Acknowledgments

We are grateful to Carlos Varela for his field assistance, Hugo Ballesteros for
lending us his collection of Chiloé frogs, and Jorge Artigas (Museo de Zoologia,
Universidad de Concepcion) for making specimens available for our study. Alice
Grandinson (British Museum) kindly examined for us the type of Cacotus cal-
caratus and Sonia Lacrampe typed the manuscript.

Literature Cited

Boulenger, C. A. 1882. Catalogue of the Batrachia Salientia of the British Museum.—London. Pp.
XVI-945.

Capurro, L. F. 1958. Lista preliminar de los Anfibios de Chile y breves apuntes sobre su distribucion

y biologia.—Investigaciones Zooldgicas Chilenas 4:289-299.

1963. Eupsophus grayi de la Isla Mocha.—Investigaciones Zooldgicas Chilenas 10:5—18.
Cei, J. M. 1958. Las laminas originales del Suplemento a los Batracios chilenos de Philippi: primera
impresion y comentarios.—Investigaciones Zoologicas Chilenas 4:265—288.

1960. A survey of the leptodactylid frogs, genus Eupsophus in Chile. Breviora 118:1-13.

1962a. Batracios de Chile.—Universidad de Chile, Santiago. 128 pp. |

1962b. El género Eupsophus en Chile.—Investigaciones Zoologicas Chilenas 8:7—42.
Formas, J. R. 1978. A new species of leptodactylid frogs (Eupsophus) from the Coastal Range in
Southern Chile.—Studies on Neotropical Fauna and Environment 13:1-9.

1980. The chromosomes of E. calcaratus and the karyological evolution of the genus
Eupsophus (Anura: Leptodactylidae).—Experientia 36: 1163-1164.

Grandinson, A. G. C. 1961. Chilean species of the genus Eupsophus (Anura; Leptodactylidae).—
Bulletin of the British Museum (Natural History) 9:111-149.

Gunther, A. 1881. III. Reptiles, batrachians, and fishes [collected during the survey of H.M.S.
‘Alert’ in the Straits of Magellan and on the coast of Patagonia].—Proceedings of the Zoological
Society of London 1881:18-22.

Lynch, J.D. 1971. Evolutionary relationships, osteology, and zoogeography of leptodactyloid frogs.—

University of Kansas, Museum of Natural History Miscellaneous Publications 53:1—238.

1972. Generic partitioning of the South American leptodactylid frog Eupsophus Fitzinger,
1843 (sensu lato).—Bulletin of the Southern California Academy of Sciences 71:2-11.

. 1978. A re-assessment of the Telmatobiinae leptodactylid frogs of Patagonia.—Occasional

Papers of the Museum of Natural History, University of Kansas 72:1—57.

Philippi, R. A. 1902. Suplemento a los Batraquios chilenos descritos en la Historia Fisica y Politica
de Chile de don Claudio Gay.—Santiago. 1-161 pp.

Instituto de Zoologia, Universidad Austral de Chile, Casilla 567, Valdivia, Chile.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 602-609

MORPHOLOGY AND DEVELOPMENT OF PLANKTONIC
LOLLIGUNCULA BREVIS (CEPHALOPODA: MYOPSIDA)

Michael Vecchione

Abstract.—Planktonic Lolliguncula brevis can be identified by chromatophore
patterns. The most consistent character in specimens <15 mm dorsal mantle
length (DML) is a cluster of three dark (‘‘red’’) chromatophores posterior to the
eyeball on the ventral surface of the head. The ventral surfaces of the mantle and
cephalic appendages are covered with numerous red chromatophores. On all
dorsal surfaces, such chromatophores are absent in small specimens (<3 mm
DML) and sparse in larger specimens.

The genus Lolliguncula is unique among cephalopods in its ability to tolerate
low-salinity waters (Brakoniecki 1980). The thumbstall squid, Lolliguncula brevis
(Blainville 1823), is a common inshore species in the Gulf of Mexico (Hixon ef
al. 1980) and along western Atlantic coasts from Delaware (Haefner 1964, Whit-
aker 1980) to the Rio de la Plata (Voss 1956). The ecology of specimens large
enough to be collected in otter trawls has been investigated (Gunter 1950, Dra-
govich and Kelly 1963, 1967, Hendrix 1979, Hixon 1980). The early life history
of this species, however, is virtually unknown.

While investigating the community ecology of planktonic molluscs in the Mid-
dle Atlantic Bight (Vecchione 1979), I found inshore loliginid specimens with
chromatophore patterns unlike those of the other common loliginid, Loligo pealei.
I speculated that these specimens were L. brevis based on their distribution and
the suggestions of Hall (1970) and Hunter and Simon (1975) that chromatophore
patterns could be used to separate species of young loliginids. Shortly thereafter,
McConathy et al. (1980) demonstrated species-specific differences in chromato-
phore patterns among hatchling loliginids with known parentage. They suggested
that chromatophore pattern differences combined with potential morphometric
differences could be used for taxonomic differentiation of young squids collected
in the field.

Squids are very difficult to rear (Yang et al. 1980). Therefore, the only practical
means of describing their development is by assembling a growth series from field
collections. I present in this report a description of the post-hatchling develop-
ment of L. brevis in the planktonic size range (<15 mm dorsal mantle length,
DML) based on specimens thus collected.

Materials and Methods

The specimens reported here have been assembled from several zooplankton
surveys. Details of collecting methods are available elsewhere (Grant 1979, Grant
and Olney 1979, Meyer 1981, Vecchione eft al. 1982). A total of 241 specimens
(size range 1.4—40.0 mm DML) were collected in ~2000 plankton tows with 153-
505 «wm mesh nets. Gear types used included bongo frames, neuston frames, pull
sleds, pushnets, and half-meter nets. All specimens were preserved in 24% form-

VOLUME 95, NUMBER 3 603

Table 1.—Specimens examined.

Dorsal mantle length
No. of

Region specimens Min. x S.D. Max.
Chesapeake Bay 15 led 4.0 5.6 23.0
Virginia shelf 49 1.8 3.6 oD) 40.0
Calcasieu estuary Da, 1.6 Doss) 0.6 4.1
Louisiana shelf 150 1.4 Mi 1.4 37

aldehyde in seawater buffered with sodium borate. Areas sampled included a
major estuary and the continental shelf of the southern Middle Atlantic Bight and
a large estuary and the continental shelf in the northwestern Gulf of Mexico (Ta-
loll 1).

All specimens were examined and measured under a dark-field dissecting mi-
croscope with ocular micrometer. Morphometric measurements were the same
as those of Vecchione (1981). Measurements were to the nearest 0.02 mm but
have been rounded to 0.1 mm because I lack confidence in the precision of
measurement of nonrigid structures to 10°? mm.

I paid particular attention to chromatophore glint. McConathy et al. (1980)
divided chromatophores into two categories: “‘reds’’ and “‘yellows.’’ I have fol-
lowed this classification and terminology. The chromatophores were usually ex-
panded on the preserved specimens analyzed in this study. This contrasted with
the typical condition of contracted chromatophores in live specimens as described
by McConathy et al. (1980). Expanded red and yellow chromatophores over-
lapped considerably. The latter were very lightly pigmented (perhaps because of
bleaching by the preservative), making it difficult to distinguish the positions of
all yellow chromatophores. I have therefore based the descriptions herein pri-
marily on the red chromatophores. The figures were drawn with expanded red
chromatophores as the planktologist would normally see them in preserved spec-

Table 2.—Morphometrics of specimens from one Middle Atlantic Bight sample. DML, dorsal man-
tle length; MW, mantle width; HL, head length; HW, head width; FL, fin length; WAF, width across
fins; AL, length of third pair of arms; TL, tentacle length.

Specimen DML MW HL HW FL WAF AL TL
1 8.9 4.4 3.0 4.1 2.6 5.4 3.0 4.4
2 Ie 3.5 al B30) Le 3.6 2.5 3.4
3 4.4 DD ID 2:2 1.0 2.5 ‘lai 1.9
4 2.6 lea 1.3 1.5 0.4 1 0.8 0.9
5 2.9 hea 1.5 (1.6 0.5 1.5 0.7 1.1
6 27 1.5 1.4 (1.5 0.5 1.4 0.3 0.8
7 2) 1.9 1.5 ihe 0.7 1.6 0.8 il. il
8 3.8 2.3 1.8 20 0.7 1.8 1.0 1a
9 2.0 13 1.1 iNet 0.4 0.9 0.3 0.8

10 Dai 2.0 1.6 ile 0.5 1.3 0.7 2
11 2.6 ew) 1.4 1.4 0.5 15 0.7 ihe
12 4.1 Del 1.9 DI 1.0 2.0 1.0 2.0
13 3.3 1.8 a7 1.7 0.6 1.8 0.8 1.3
14 a is 1.6 17 0.4 1.4 0.6 0.9

604 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Lolliguncula brevis (DML = 2.3 mm) showing placement of red chromatophores; left,
ventral view; right, dorsal view. Scale line = 2 mm.

imens. Because few specimens were >4.5 mm DML and because many specimens
of all sizes appeared damaged during collection or preservation, the following
descriptions were based on specimens which were arbitrarily selected based on
state of preservation within representative size ranges.

Identification and Development

The smallest specimens considered in this study closely matched the descrip-
tion by McConathy et al. (1980) of L. brevis hatchlings. For example, a 2.3 mm
DML specimen (Fig. 1) lacked dorsal red chromatophores on either the head or
the mantle. On the ventral surface of the head posterior to each eyeball, a tri-
angular cluster of 3 red chromatophores matched that described for L. brevis.
The number of red chromatophores on the ventral mantle surface (43) was within
the range for hatchling L. brevis (36-44) but well above those of Loligo pealei
and L. plei (15-32). Mantle chromatophores, however, were not arranged in the
7 transverse rows described in L. brevis by McConathy et al. (1980). Red chro-

VOLUME 95, NUMBER 3 605

"ey,
ot

oe

‘4.
bo
te
‘t
o
s

"s

f

a
ce

Fig. 2. Lolliguncula brevis (DML = 4.5 mm) showing placement of red chromatophores; left,
ventral view; right, dorsal view. Scale line = 2 mm.

matophores on the cephalic appendages were confined to the ventral surfaces of
the tentacles and fourth pair of arms. Yellow chromatophores were discernable
on this specimen at the approximate locations described by McConathy et al.
(1980).

A 4.5mm DML specimen of L. brevis, representative of the 3.5—5.5 mm DML
size range, had red dorsal chromatophores, 4 on the mantle and 6 on the head

606 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

<p

GOO OE
* Sage
@ @

Fig. 3. Lolliguncula brevis (DML = 9.7 mm) showing placement of red chromatophores; left,
ventral view; right, dorsal view. Scale line = 2 mm.

(Fig. 2). I have not yet found any specimens with only 2 red chromatophores on
the dorsal mantle. I therefore cannot state the size at which these features first
develop. Ventral placement of red chromatophores was similar to that of smaller
specimens, including triangular clusters posterior to the eyeballs and numerous
chromatophores (50) on the ventral mantle. Chromatophores had been added to
the ventral rows on the tentacles and fourth arms. A second row of very dark
chromatophores had also been added to the tentacles at the bases of the ventral
suckers. This row was present in all specimens >4.5 mm DML but absent in
smaller specimens. At 4.5 mm DML the third pair of arms had developed a

VOLUME 95, NUMBER 3 607

swimming margin. The fins remained separated at the posterior mantle extremity.
Three pronounced anterior mantle lobes were present in this specimen, one dorsal
and 2 ventrolateral. Similar but smaller lobes were seen in a 3.8 mm DML spec-
imen but were absent in smaller specimens.

Variability in chromatophore development was apparent in 2 specimens of 6.6
mm DML and one of 7.1mm DML. All of these specimens were similar in overall
morphology to the 4.5 mm DML specimen described above. Numbers of mantle
chromatophores varied, however, as follows:

dorsal- ventral-

mantle mantle

DML chrom. chrom.
6.6 12 52
6.6 15 47
ea 9 48

In a 7.4 mm DML specimen the fins were joined just posterior to the mantle end.
This specimen had 14 dorsal- and 51 ventral-mantle chromatophores.

The specimen in Fig. 3 measured 9.7 mm DML. Gross morphology included
large anterior dorsal and ventrolateral mantle lobes, fins joined posterior to the
mantle end, and comparatively large eyeballs. Triangular clusters of red chro-
-matophores were located ventroposterior to the eyeballs. A third row of chro-
matophores on the tentacles was located on the recently developed swimming
margins. The swimming margins on the third pair of arms had developed a row
of chromatophores. Red chromatophores on the mantle included 21 dorsal and
75 ventral. The dorsal surface of the head contained 8 red chromatophores and
the second pair of arms each had 2 red dorsal chromatophores.

A 13.7 mm specimen was similar in appearance to the juvenile L. brevis com-
monly taken in otter trawls. There were 33 dorsal and 98 ventral red chromato-
phores on the mantle. The number of chromatophores indicated rapid increase
of chromatophores in this size range. The dorsal surface of the fins, which were
joined posterior to the mantle, had numerous small red chromatophores along
the posterior edge as well as small yellow chromatophores scattered over the
entire dorsal surface.

A limited set of morphometric observations on specimens 2.0-8.9 mm DML
from a single sample (Table 2) were not significantly different from similar ob-
servations on Loligo pealei described by Vecchione (1981).

Discussion and Conclusions

Morphometric indices are not suitable for separating the young of the two
common genera of inshore squids. However, given fresh specimens it should be
possible to separate L. brevis from L. pealei and L. plei using differences in the
ratio of dorsal : ventral red mantle chromatophores at a given size. This character,
though, is increasingly variable in larger (>5 mm DML) specimens.

The size-frequency distribution of planktonic specimens was strongly skewed
toward small sizes. These small individuals (<4 mm DML) were identifiable by
use of the chromatophore criteria of McConathy et al. (1980). Most lacked red
dorsal chromatophores and all had high numbers of red chromatophores on the

608 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ventral surface of the mantle. These ventral chromatophores were not, however,
organized into 7 distinct transverse rows as described by McConathy et al. (1980).

Figures of late-stage embryos of L. brevis by Hall (1970) and Hunter and Simon
(1975) show chromatophore patterns which do not match those presented here.
I agree, however, with McConathy et al. (1980:279) that ““Because the primary
interest of these studies was either morphological or embryological and not spe-
cifically about chromatophores, the chromatophore arrangements depicted may
be inexact.”’

Chromatophores on the mantle are subject to abrasion in net-captured speci-
mens. Head chromatophores seem to be more protected, perhaps because the
head can be partly withdrawn into the mantle. Of the specimens examined for
this study, all but one had 3, rather than 4, red chromatophores posterior to the
eyeballs on the ventral surface of the head. The one exception appeared to have
been damaged as it had 3 uniform-sized chromatophores on one side and 2 similar-
sized and 2 small (fragments?) chromatophores on the other side. This triangular
cluster is a very consistent character suitable for generic identification of plank-
tonic specimens, contrasting with the condition in L. pealei and L. plei of a
trapezoidal arrangement of 4 red chromatophores (McConathy et al. 1980; Vec-
chione, pers. obs.).

Acknowledgments

It is a pleasure to acknowledge the assistance of Cathy Meyer and Don Weston,
whose comments improved an earlier draft of this paper, and of Deanne Bonvil-
lain, who typed the manuscript. Comments by an anonymous reviewer were very
helpful in finalizing the manuscript. Contribution No. 1071 of the Virginia Institute
of Marine Science.

Literature Cited

Brakoniecki, T. F. 1980. Lolliguncula tydeus, a new species of squid (Cephalopoda; Myopsida)
from the Pacific coast of Central America.—Bulletin of Marine Science 30:424—-430.

Dragovich, A., and J. A. Kelly. 1963. A biological study and some economic aspects of squid in

Tampa Bay, Florida. Pp 87—102 in Gulf and Caribbean Fisheries Institute, Proceedings of the

15th Annual Session.

, and 1967. Occurrence of the squid, Lolliguncula brevis in some coastal waters of

western Florida.—Bulletin of Marine Science 17:840-844.

Grant, G. C. 1979. Middle Atlantic Bight zooplankton: Second year results and a discussion of the

two-year BLM-VIMS survey.—Virginia Institute of Marine Science Special Report in Applied

Marine Science and Ocean Engineering 192, 236 pp.

, and J. E. Olney. 1979. Lower Bay zooplankton monitoring program: An introduction to

the program and results of the initial survey of March 1978.—Virginia Institute of Marine

Science Special Scientific Report 93, 92 pp.

Gunter, G. 1950. Seasonal population changes and distributions as related to salinity, of certain
invertebrates of the Texas coast, including commercial shrimp.—Publications of The Institute
of Marine Science 1:7-51. :

Haefner, P. A. 1964. Morphometry of the common Atlantic squid, Loligo Pealei, and the brief
squid, Lolliguncula Brevis in Delaware Bay.—Chesapeake Science 5:138-144.

Hall, J. R. 1970. Description of egg capsules and embryos of the squid, Lolliguncula brevis, from
Tampa Bay, Florida.—Bulletin of Marine Science 20:762-768.

Hendrix, J. P. 1979. Osmotic concentration changes in the bay squid Lolliguncula brevis (Blainville,
1823) at various salinities.—Bulletin of the American Malacological Union 1979:69-70.

VOLUME 95, NUMBER 3 609

Hixon, R. F. 1980. Growth, reproductive biology, distribution and abundance of three loliginid

squid species (Myopsida, Cephalopoda) in the Northwest Gulf of Mexico.—Ph.D. dissertation,

University of Miami, 92 pp.

, R. T. Hanlon, S. M. Gillespie, and W. L. Griffin. 1980. Squid fishery in Texas: Biological,

economic, and market considerations.—Marine Fisheries Review 42:44—S0.

Hunter, V. D., and J. L. Simon. 1975. Post-cleavage morphology in the squid Lolliguncula brevis
(Blainville, 1823).—Veliger 18:44—51.

McConathy, D. A., R. T. Hanlon, and R. F. Hixon. 1980. Chromatophore arrangements of hatchling
loliginid squids (Cephalopoda, Myopsida).—Malacologia 19:279-288.

Meyer, C. E. 1982. Zooplankton communities in Chesapeake Bay seagrass systems.—M.S. Thesis,
College of William and Mary, 88 pp.

Vecchione, M. 1979. - Planktonic molluscan faunal structure across a large scale environmental

gradient.—Ph.D. dissertation, College of William and Mary, 153 pp.

. 1981. Aspects of the early life history of Loligo pealei. Journal of Shellfish Research 1:171—180.

——, C. E. Meyer, and C. L. Stubblefield. 1982. Zooplankton. Chapter 8 in Final Report on
Contract DE-AC-96-80P010228 with the U.S. Department of Energy. McNeese State Univer-
sity, Lake Charles, Louisiana. 69 pp.

Voss, G. L. 1956. A review of the cephalopods of the Gulf of Mexico.—Bulletin of Marine Science
of the Gulf and Caribbean 6:85-178.

Whitaker, J.D. 1980. Squid catches resulting from trawl surveys off the southeastern United States.—
Marine Fisheries Review 42:39-43.

Yang, Y. T., R. T. Hanlon, R. F. Hixon, and W. H. Hulet. 1980. First success in rearing hatchlings
of the squid Loligo pealei Lesueur 1821.—Malacological Review 13:79-80.

Research and Development, McNeese State University, Lake Charles, Loui-
-siana 70609.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 610-620 -

REDESCRIPTION OF EXOEDICEROS FOSSOR (STIMPSON,
1856) AN AUSTRALIAN MARINE FOSSORIAL
AMPHIPOD, THE TYPE-GENUS OF THE
NEW FAMILY EXOEDICEROTIDAE

J. Laurens Barnard and Margaret M. Drummond

Abstract.—The type-species of Exoediceros, for which the type material has
been lost, is redescribed from fresh Australian material and compared to its junior
synonym Exoediceros arenicola (Haswell), for which type-material is available.
This is compared to Exoediceros maculosus Sheard (1936), a sympatriot. A new
family, Exoedicerotidae, is described to include Exoediceros, Exoediceropsis,
Bathyporeiapus, Metoediceros, Parhalimedon, and Patuki, all southern 2-eyed
(or blind) genera with apical spination on the rami of uropods 1-2. These are
assumed to be more primitive than the advanced Oedicerotidae which have dom-
inated the northern hemisphere and the deep-seas.

Several genera formerly identified as Oedicerotidae, Exoediceros, Exoedice-
ropsis, Bathyporeiapus, Metoediceros, Parhalimedon, and Patuki, are now con-
sidered to be distinguishable as a new family to stand in a position plesiomorphic
to the Oedicerotidae. The type-species of the type-genus, Exoediceros, is rede-
scribed herein so as to establish this family.

Legends: Capital letters describe morphological parts; lower case letters to left
of capital letters denote specimens cited in figure legends, lower case letters to
right of capital letters or in body of drawing are cited in following list: A, antenna;
B, body; C, coxa; D, dactyl; E, pleon; F, accessory flagellum; G, gnathopod; H,
head; I, inner plate or ramus; J, gill; K, palp; L, labium; M, mandible; N, pleopod;
O, outer plate or ramus; P, pereopod; Q, brood plate; R, uropod; S, maxilliped;
T, telson; U, prebuccal anterior; X, maxilla; Y, calceolus; Z, aesthetasc; m,
medial; r, right; s, setae removed.

Exoedicerotidae, new family

Diagnosis.—Amphipoda-like Oedicerotidae but apices of rami on uropods 1-2
spinose; eyes, when present, paired.

Description.—Body laterally compressed, scarcely or not processiferous, uro-
somites 1-3 free. Head not strongly galeate, rostrum medium, thin or weak.

Peduncles of antennae medium to short, flagella usually calceoliferous, calceoli
probably of oedicerotid kind 7 (Lincoln and Hurley 1981), accessory flagellum 0-
l-articulate.

Labrum weakly excavate or entire, epistome occasionally produced. Mandible
strong, incisor toothed, raker row well developed, molar triturative to obsoles-
cent, palp 0-3-articulate. Inner lobes of lower lip fleshy and separate. Maxillae
well developed, inner plates poorly to strongly setose. Maxillipeds well devel-
oped, outer plate small to large, palp 4-articulate.

Coxae 1-4 well developed, coxa 4 lobate and excavate or not, coxa 5 generally

VOLUME 95, NUMBER 3 611

only slightly smaller than 4. Gnathopods diverse, ranging from medium, alike and
subchelate with spine fields on hands, to feeble, simple and elongate and without
spine fields. Pereopods 3--7 fossorial, with powerful and spinose articles, dactyls
often vestigial or absent on pereopods 3-6, pereopod 7 very large as in Oedicerot-
idae, article 2 broadly expanded, appendage much longer than pereopods 5-6,
dactyl elongate and well armed.

Pleon powerful, pleopods well developed, epimera ordinary, poorly ornament-
ed. Uropods 1-2 with long rami bearing apical spines. Uropod 3 variable, mod-
erately to strongly developed or vestigial. Telson short, entire, laminar.

Gills on coxae 2-7; oostegites narrow to moderately broad.

Exoediceros Stebbing

Exoediceros Stebbing, 1899:208 (Oedicerus fossor Stimpson, 1856, original des-
ignation).

Diagnosis.—Body not carinate. Eyes paired, separate. Article 3 of peduncle
of antenna 1 half or less as long as article |. Fully articulate, scale-like accessory
flagellum present. Primary flagellum of antenna | with similar articles bearing
similar armaments. No articles of antenna 2 especially swollen. Mandibular in-
cisor projecting, toothed; molar large, triturative; palp 3-articulate, article 2 straight,
article 3 clavate, stubby. Inner lobes of lower lip distinct, separate, fleshy. Outer
plate of maxilla 2 lacking thick spine. Plates of maxilla 2 diverse. Coxae poorly
' setose, coxae 1-4 rounded below, coxa 4 subrectangular, scarcely excavate pos-
teriorly, not lobate. Gnathopods sexually dimorphic, in each sex similar to each
other, subchelate, wrists weakly lobate, not guarding hands, palms oblique, hands
with dense fields of blunt spines near apex of closed dactyl. Dactyl of pereopods
3-4 obsolescent. Coxal gill 5 minute. Article 2 of pereopod 7 expanded but scarce-
ly lobate. Uropod 2 not reaching far along uropod 3; peduncle of uropod 3 scarcely
elongate, with large marginal spines, rami short. Telson entire.

Relationship.—TYhe apparent closest relative of this genus is Metoediceros
Schellenberg (1931) which has similar gnathopods bearing spine fields. Meto-
ediceros differs from Exoediceros in the reduction of uropod 3 to a small vestige,
the lack of an accessory flagellum, the lack of a mandibular palp and the poorly
setose inner plate of maxilla 1.

Exoediceropsis Schellenberg (1931) differs from Exoediceros in: (1) the feeble
molar; (2) the feeble, mitten-shaped gnathopods; (3) the lobate coxa 4; (4) the
nonsetose inner plate of maxilla 1; and (5) the small outer plate of the maxilliped.

Bathyporeiapus Schellenberg (1931) differs from Exoediceros in items 1, 2 and
4 as cited above. Parhalimedon Chevreux (1906) differs from Exoediceros in
items | and 2 above plus (6) the long uropod 3 with long peduncle; (7) the poorly
armed rami of uropod 3; and (8) the absence of eyes.

The male of Patuki Cooper and Fincham (1974) is unknown but the uropod 3
of the female is shorter than uropods 1 and 2, with unarmed peduncle; and the
eyes are closer together dorsally than those of Exoediceros.

Exoediceros fossor (Stimpson)
Figss 5

Oedicerus fossor Stimpson, 1856:394.—Bate, 1862:373.—Haswell, 1882:238.
Oediceros fossor.—Della Valle, 1893:556.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

612

{7
ly Ly

SS We a
Y a:
LM isege

Me ifaee eseaee Ae 7 \
A
—~ d y
ane oe,
ALIN iD

YAf Azm

Fig. 1. Exoediceros fossor, male ‘‘p”’ 8.01 mm.

VOLUME 95, NUMBER 3 613

Exoediceros fossor.—Stebbing, 1906:239; 1910:638.
Oedicerus arenicola Haswell, 1879:325, pl. 24, fig. 3; 1882:239-240.—Della Valle,
1893:556.

Diagnosis.—Rostrum short, not reaching beyond middle of article | of antenna
1. Plates of maxilla 2 broad. Coxa | narrow and tapering distally. In life body
said to lack chromatophores (Sheard 1936).

Description of male ‘‘p’’.—Each eye heavily pigmented. Ommatidia clear api-
cally. Lateral cephalic lobes small, mammilliform.

Antennae short, extending subequally, articles of flagella short, bead-like, pro-
liferate, on antenna | basal articles with | large calceolus of oedicerotid kind 7
(Lincoln and Hurley 1981), each basal article also with small aesthetasc of same
but rudimentary kind situated on outer face of article obliquely proximal to main
calceolus then towards apex each article tending to bear 2—3 small versions of
calceoli, all articles also with 2 aesthetascs each; formula of calceoli (L = large,
s = small) on flagellum of antenna 1 = 0,s,2s,sL,LL,Lss,Lss,Lss,Lss,Lss,
SS,SSS,SSS,SSS,SSS,SS,SS,S ... broken (probably only final article miss-
ing); formula on antenna 2 = s,LL,LLs,LL,LL,LL,LL,LLs,Lss,Lss,
Lss,sss,sss,sss,s,0; no clavate aesthetascs on antenna 2.

Upper lip with tiny ventral notch. Incisors toothed; right lacinia mobilis
3-pronged, prongs serrate; left lacinia mobilis with 5 teeth; rakers stout, right and
left about 9 each; molar stout, cuboid but moderately triturative; palp stout,
article | short, article 2 expanded and strongly setose, article 3 clavate, setae =
ABDE. Inner plate of maxilla 1 fully setose medially; outer plate with 11 spines
(not all shown on illustration); palp strongly setose, 2-articulate. Plates of maxilla
2 broad, inner with full oblique facial row of setae. Inner plates of maxilliped
with medial margins appressed and bent orally, setose, apices each with 2 small
medial spines and numerous widely spread setae; outer plates not larger than
inner, medially spinose; dactyl unguiform, with small apical nail and several se-
tules on inner margin.

Coxa 5 scarcely shorter than coxa 4. Gnathopod 2 slightly larger than 1, both
weakly twisted in death. Dactyls of pereopods 3-4 extremely minute, each bear-
ing ordinary setule itself remaining normally large and thereby dwarfing dactyl.
Pereopods 5-6 bearing small dactyls with largely absorbed apical nail and large
setule. Gills present on coxae 2-7, flat, unpleated, with transverse capillaries,
gills of coxae 2-3 ovate, sac-like, of coxa 4 adz-shaped, of coxa 5 very small,
tear-drop shaped and pediculate, of coxa 6 larger, ovate, of coxa 7 larger than 5,
like dried leaf with base twisted into brood space.

Pleopods relatively similar, peduncles elongate, each with 2 feeble retinacula,
each outer ramus with posterior tooth or boss on article 1, outer and inner rami
about 1.5 and 1.3 times respectively as long as peduncles, outer and inner rami
with about 19 and 15 articles respectively. Epimera 1-3 each with several an-
teroventral marginal setae, epimeron | with distinct facial ridge bearing several
spinules, ridge of epimeron 2 with spinule row disjunct above and epimeron 3
with facial ridge but no spines; posteroventral corners of epimera 1—3 rounded.

Urosomite 1 with 2 weak dorsal humps, urosomites 2—3 each with sharp pos-
terodorsal edge, urosomite 3 so high as to obscure most of telson from lateral
view. Peduncle of uropod | with basofacial row of setules and spinule, dorsolat-

614 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Exoediceros fossor, male ‘‘p’’ 8.01 mm.

VOLUME 95, NUMBER 3 615

eral margin naked except for several basal setules, medial margin with 4 medium
spines in widely disjunct tandem, peduncle of uropod 2 with 2 widely spread
dorsal spines, | apicomedial spine; rami of uropods 1-2 all with 2 apical nails and
2 minute subapical accessory nails, but accessory nail on inner ramus of uropod
2 vestigial; inner rami otherwise naked on dorsal margins, outer rami of uropods
1-2 with 2 and 1 dorsal spines respectively. Peduncle of uropod 3 with 3 dorso-
lateral spines, 2 dorsomedial spines and 2 spines and 2 setules in tandem, rami
shorter than peduncle, weakly foliate, apically and medially setose, outer ramus
with subbasal ridge bearing terminal spine, inner ramus with subbasal medial
spine. Telson very short, apex rounded, subtruncate, each dorsolateral face with
2 pairs of pencillate setules from about M. 50 to M. 80. Cuticle very minutely
punctate.

Male ‘‘n’’.—Spine count on epimeron 2 = 8-4-2; dorsolateral margin of pe-
duncle on uropod 2 with 3 spines.

Male ‘‘c’’, smallest available male.—Like adult but flagellum of antenna 1| with
calceolus formula of 0-1-2-2-2-2-2-2-1 (broken), aesthetasc formula = 1-1-1-2-2-2-
2-2-1, calceolus formula of flagellum on antenna 2 = 0-1-2-2-3-3-3-2 (broken), all
calceoli small on both pairs of antennae. Gnathopods lacking medial fields of
spines on faces of hands, gnathopod | with defining spine on each side of hand
followed behind by 2 spines in tandem on each side, gnathopod 2 with same
scheme but 3 following spines on each side. Facial formula of setae on epimeron
2 = 2-1-1. Peduncle of uropod | with 2 basofacial spines, 1 dorsolateral spine on
outer ramus; peduncle of uropod 2 with 2 dorsal spines, neither ramus with dorsal
spine; peduncle of uropod 3 with 1 dorsal spinule, 1 apical blunt spine. Cuticle
grossly scalloped.

Female ‘‘i’’.—Differing from male in presence of more small calceoli on an-
tennal flagella but absence of the large variety; formula of small calceoli on fla-
gellum of antenna | = 0,1,2,1,3,4,4,4,5,4,4,4,5,5,4,3 . . . (broken, probably only
last article missing), only 1 aesthetasc per article; formula of small calceoli on
flagellum of antenna 2 = 0,1,2,3,4,4,4,4,5,5,5,5,5,5, 5. . . (broken, probably only
last article missing), no aesthetascs present.

Gnathopods, especially hands, much smaller than in male, lobes of wrists much
broader, hands more evenly ovate, palms weakly convex, no spine fields present.

Brood plates thin, strongly setose (one illustrated), pair of coxa 5 half as long
as Other 3 pairs but as broad.

Uropod 3 and telson as in male.

Other minor differences not sexually related: spine count on face of epimeron
2 = 5-2-2-1; setae on inner ramus of uropod 3 = 11-12, outer = 12; left inner
ramus also with 2 spines (not | as in male), peduncles of right and left sides with
2 and 3 spines each.

Juvenile ‘‘j’’ 2.16 mm.—Flagellum of antenna | with 5 articles, calceolus for-
mula = 0-0-1-0-0, aesthetasc formula = 0-2-1-1-0; calceolus formula on flagellum
of antenna 2 = 0-0-1-0, all calceoli small. Dactyl of pereopods 3-4 no larger than
in adults relative to appendages. Formula of setae on epimeron 2 = 2-0-0. Pe-
duncle of uropod 1 with 1 apicolateral long thin spinule, rami lacking dorsal
Spines, apex of outer ramus with 2 spines and 2 large subapical spines, inner with
2 apical, 1 subapical large and 1 tiny subapical spinule. Peduncle of uropod 2 with
1 spine on each apicodorsal corner, rami lacking dorsal spines, apex of outer

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VOLUME 95, NUMBER 3 617

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mm.

ramus with 2 large spines, | large and | tiny subapical spines, inner ramus with
2 large apical, | large and no other subapical spine. Peduncle of uropod 3 with 1
Spine on each apicodorsal corner, inner ramus with 1 thin dorsomarginal spine
and 2 apical setae, outer ramus without spine, with 3 apical and 0-1 (right or left)

618 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

7
DD An

a
=

Fig. 5. Exoediceros fossor, unattributed figures = male ‘‘p’’ 8.01 mm; i to left = female ‘‘1’’ 7.49
mm; r to left = female ‘‘r’’ 7.09 mm.

VOLUME 95, NUMBER 3 619

subapical seta. Apical pair of setules on telson almost at posterior margin. Cuticle
with large scallops.

This scallop pattern, present in the cuticle of all juveniles and small specimens
examined, is the result of an orderly arrangement in arcs of the minute, rounded,
pebble-like bodies in it (as in E. maculosus). This scalloping is not apparent in
large individuals where the cuticular bodies are more evenly distributed in a flat
pattern of pentagons or hexagons (themselves sometimes ill defined).

Type-locality.—Australia, Botany Bay (fossor); Shark Island, Port Jackson (ar-
enicola). Types of fossor probably lost in Chicago Fire of 1871; probably no types
of arenicola ever selected: we hereby select as lectotype male ‘‘a’’ 8.42 mm,
from New South Wales Museum no. 10406, Port Jackson, New South Wales,
assumed to be the original material of Haswell; also accompanying this male is
a female *‘b’’ 7.25 mm.

Voucher material.—Towra Point, New South Wales, intertidal sand, 23 August
1980, coll. Dr. Deborah Dexter, male ‘“‘p’’ 8.01 mm (main illustration), female
‘‘7’’ 7.49 mm (main female described and illustrated), male ‘‘n’’ 6.55 mm, female
‘“‘q’’ 6.63 mm, female ‘‘r’’ 7.09 mm (illustrated); Mallacoota, Victoria, 9 February
1978, still water, intertidal, coll. M. M. Drummond, smallest available male ‘“‘c’’
3.40 mm (described); Gippsland Lakes, Victoria, at sand spit east of Lakes En-
trance, 1 April 1976, coll. P. Hutchings and J. D. Kudenov, juvenile ‘‘j’’ 2.16 mm
(described and illustrated), young male ‘‘k’’ 5.71 mm, young female ‘“‘y’’ 4.50

mm.

Additional material.—NSW: Port Jackson, JKL Australian Museum (3); Towra
Point, Botany Bay, D. Dexter (100+); Narabeen, DD (3); Careena Bay, st. 68,
Georges River, EBS (4); Merimbula, J. H. Day Sample 2B, 9 May 1975 (100+);
Merimbula, MMD samples Feb. 1972—Dec. 1978 (100+). Victoria: Mallacoota, J.
D. Kudenov (20), MMD, 9 Feb 1978 (40); Tidal River, Wilsons Promontory,
MMD, 31 Oct. 1978 (35); Gippsland Lakes, Lakes Entrance, P. H. and J. D. K.,
April 1976, 2 samples from 2 stations (10). Tasmania: Anson’s Bay, May 1978,
Tasmanian Fisheries Development Authority, D. Hoggins (12).

Relationship.—Exoediceros maculosus Sheard (1936) differs from E. fossor in
many characters, among them the following: (1) the long rostrum; (2) the short
article 2 of antenna 1; (3) the regular occurrence, on antennal flagella, of swollen
articles, alternating with ordinary articles in a ratio either of 1:1 or, particularly
in the middle section, 1:2, on male antennae only these swollen articles bearing
large calceoli and a battery of 4—5 simple aesthetascs, alternating articles with
small calceoli or none, swollen articles on female antenna 1 bearing aesthetascs
in addition to small calceoli; (4) the long, straight blade of the mandibular incisor,
with teeth confined to either end; (5) the leaf-like semifalcate article 3 of the
mandibular palp, longer than article 2; (6) the 4-cusped right lacinia mobilis; (7)
the lack of strong distinction between male and female gnathopods, those of the
male bearing no medial spine fields but 3 more or less, seriate ranks of spines;
(8) the short gnathopodal article 5 in both sexes; (9) the rudimentary dactyl on
pereopods 3 and 4; (10) the regular and even setation on epimeron | in females;
(11) the regular and even facial spination (not setation) on epimeron 2; this epi-
meron with midvertical facial ridge; (12) regular facial spination of epimeron 3;
(13) regular, even spination of peduncle and rami on uropods 1 and 2; (14) short-
ened outer ramus of uropod 2; (15) relatively short peduncle of uropod 3 (shorter
than rami); (16) the apically 2-notched telson.

620 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

A separate paper on Exoediceros maculosus will be published elsewhere. Prob-
ably Oedicerus latrans Haswell (1879) is a senior synonym of E. maculosus.

Distribution.—Port Jackson, New South Wales to southeastern Victoria, and
Tasmania, protected beaches, intertidal or shallow sands.

Acknowledgments

We thank Deborah Feher for inking our drawings and Dr. Thomas E. Bowman
for his advice. We also gratefully acknowledge the help of colleagues who made
available material: Dr. J. K. Lowry, The Australian Museum; Dr. G. C. B. Poore,
National Museum of Victoria (MSG Victoria Gippsland Lakes Survey); Dr. Philip
Gibbs, NSW Fisheries Department (NSW Fisheries Estuarine Benthic Survey,
EBS); D. D. Hoggins, Tasmanian Fisheries Development Authority; Professor
Emeritus J. H. Day, South Africa (Merimbula); Dr. Deborah Dexter, University
of San Diego California (beach sands collections from the environs of Sydney);
Dr. J. D. Kudenov, formerly of Marine Studies Group, Victoria (now University
of Alaska) (Mallacoota and Gippsland Lakes).

Literature Cited

Chevreux, E. 1906. Crustacés amphipodes.—Expédition Antarctique Frangaise (1903-05) Com-
mandée par le Dr Jean Charcot. Sciences Naturelles: Documents Scientifiques: 100 pp., 56
figs.

Cooper, R. D., and A. A. Fincham. 1974. New species of Haustoriidae, Phoxocephalidae, and
Oedicerotidae (Crustacea: Amphipoda) from northern and southern New Zealand.—Records
of the Dominion Museum 8:159-179, 13 figs.

Della Valle, A. 1893. Gammarini del Golfo di Napoli.mFauna und Flora des Golfes von Neapel,
Monographie 20:xi + 948 pp. 61 plates (in separate atlas).

Haswell, W. A. 1879. On some additional new genera and species of amphipodous crustaceans.—

Proceedings of the Linnean Society of New South Wales 4:319-350, plates 18-24.

1882. Catalogue of the Australian stalk- and Sessile-eyed Crustacea.—Australian Museum,

Sydney:xxiv + 324 pp., plus Addenda et Corrigenda, 4 plates.

Lincoln, R. J., and D. E. Hurley. 1981. The calceolus, a sensory structure of gammaridean am-
phipods (Amphipoda:Gammaridea).—Bulletin of the British Museum (Natural History), Zo-
ology 40:103-116, 4 figs.

Schellenberg, A. 1931. Gammariden und Caprelliden des Magellangebietes, Sudgeorgien und der
Westantarktis.—Further Zoological Results of the Swedish Antarctic Expedition 1901-1903,
2(6), 290 pp., 136 figs, 1 plate.

Sheard, K. 1936. Amphipods from a South Australian reef. Part I.—Records of the South Australian
Museum 5:445—455, 4 figs.

Stebbing, T. R. R. 1899. Revision of Amphipoda (Continued).—Annals and Magazine of Natural
History (7)4:205-211.

——. 1906. Amphipoda I. Gammaridea.—Das Tierreich 21, 806 pp., 127 figs.

1910. Crustacea. Part 5. Amphipoda.—Scientific Results of the Trawling Expedition of the

H.M.C.S. ‘‘Thetis.’’ Australian Museum, Memoir 4, 2:565—658, plates 47*—60*.

Stimpson, W. 1856. Descriptions of some new marine Invertebrata.—Proceedings of the Academy
of Natural Sciences, Philadelphia 7:385-394.

(JLB) Department of Invertebrate Zoology, Smithsonian Institution, Washing-
ton, D.C. 20560, USA; (MMD) National Museum of Victoria, Melbourne, Vic-
toria, Australia 3000.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 621-624

COURTSHIP DISPLAY IN A BORNEAN FROG
Keith A. Harding

Abstract.—An extraordinary courtship display by a ranid frog, Staurois parvus,
was observed above a waterfall on a lowland forest river in Brunei, North Borneo,
in August 1981. The display consisted of kicking out one of the hind limbs re-
vealing the pale blue webbing of the foot which contrasted strongly with the frog’s
buff and olive dorsal coloration. The display was filmed and a print of a complete
42-frame sequence is reproduced in this paper.

During July and August 1981, I was a member of a British Broadcasting Cor-
poration (BBC) Natural History Unit expedition to southeast Asia to film forest
species for a television series. The expedition camped at Labi, an area of lowland
forest on the Rampayoh River, Brunei, North Borneo from 1 to 5 August. The
river at this site was about 15 m wide and there was a waterfall about 5 m high
(Fig. 1).

On the evenings of 2 and 3 August, several small frogs (approx. 3 cm SV),
later identified as Staurois parvus Inger and Haile, were observed on the rock
surrounding the waterfall and a shrill chirping call was heard. At about 0800 hr
on 4 August, Adrian Warren drew my attention to a single frog on a ledge above
the waterfall (Fig. 2). The frog slowly and deliberately kicked out its right hind
leg exposing the pale blue webbing of the foot, which contrasted strongly with

Fig. 1—Waterfall on Rampayoh River, Brunei where frog courtship was observed. Exact spot
(Fig. 2) arrowed.

622 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2.—Rock ledge above waterfall with ¢ Staurois parvus (arrowed) about to display.

its buff and olive dorsal coloration and with the immediate habitat. This striking
display was clearly visible from the opposite bank of the river as a sudden flash
of blue. The display was repeated several times and later the same frog was
observed in amplexus with a larger (approx. 4 cm SV) female. The pair was then
lost from view in the surrounding vegetation. No calling was heard during the
display.

This behavior was filmed by Rodger Jackman with a Bolex EL 16 mm camera
on Eastmancolor 7247 negative film. A 42-frame sequence has been reproduced
from a positive rush print (Fig. 3) in strips of 7 frames (the first frame in each
strip being numbered for clarity). The film was shot at standard television speed
(25 frames/second).

The display reported here represents an undescribed form of anuran courtship
behavior. It is possible that other Bornean frogs display in a similar manner as
several species have brightly colored webbing (Inger, personal communication).
Future observations may complete the life-histories of these little-known Bornean
species.

Acknowledgments

Julian Dring and Alice Grandison of the British Museum (Natural History)
kindly identified the frog. Iam most grateful to Adrian Warren of the BBC Natural
History Unit for inviting me to join the expedition and for reading the manuscript,
and to Richard Brock, Executive Producer of ‘‘Planet Earth’’, for permission to
publish this paper. Robert F. Inger of the Field Museum of Natural History,
Chicago, kindly commented on the photographic material.

5 Drakewalls Place, Gunnislake, Cornwall, PL18 9EJ, England.

VOLUME 95, NUMBER 3 623

Fig. 3.—Courtship display of Staurois parvus (see text) a. frames 1-21; b. frames 22-42.

624 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 625-636

ANTHURIDS FROM THE HOUTMAN ABROLHOS ISLANDS,
WESTERN AUSTRALIA (CRUSTACEA:
ISOPODA: ANTHURIDAE)

Brian Kensley and Gary C. B. Poore

Abstract.—Six species of anthurid isopods, three or probably four new, are
recorded from coral and algal rubble at Rat Island in the Houtman Abrolhos
Islands. Apanthura zeewykae n. sp., Eisothistos bataviae n. sp., and Panathura
haddae n. sp. are first records of their respective genera from Western Australia.
The range of Mesanthura protei Kensley is extended from the western Indian
Ocean. Panathura ardea (Poore and Kensley), previously known from Queens-
land, is herein removed from Coralanthura. Heptanthura sp. is thought to be
undescribed.

Knowledge of the anthuridean isopods of Western Australia is confined to six
species described by Thomson (1946, 1951). A small collection made for us by
Dr. P. Hutchings at the Houtman Abrolhos Islands (28°43'S, 113°47’E) adds to
what must be a considerably larger fauna. Affinities to the fauna of other parts
of the Indian Ocean are suggested here (see Kensley 1980).

Collections were made at ten stations on the eastern side of Rat Island. Coral
and algal rubble samples were taken from the bases of dense stands of live Ac-
ropora coral on the reef crest at 1 meter depth. Tufted red and green algae
overgrew the rubble. Four samples were taken from a similar substrate on a sandy
shoal east of Rat Island in 3—5 meters depth. The samples from the reef crest
were combined, as were those from the sandy shoal. As well as the anthurids
reported on here, other isopods belonging to the genera Gnathia, Stenetrium,
Carpias, Munna, and Limnoria were recorded.

Material has been deposited in the Australian Museum, Sydney (AM), the
National Museum of Victoria, Melbourne (NMV), and the United States National
Museum of Natural History (USNM).

Apanthura zeewykae, new species
Rigs 2

Description.—Integument thin, head with transverse pigment band between
eyes; pereonites 1-3 with pigment patch at anterolateral corners, pereonites 4—7
with scattered pigmentation posteriorly; pleon with 5 pairs of dorsal pigment
patches. Body proportions: C < 1 >2>3<4=5<6.>/7. Head with trian-
gular rostrum reaching beyond anterolateral corners. Dorsolateral eyes large, well
pigmented. Pleon about 1.5 as long as pereonite 7; pleonites 1—S fused, pleonite
6 free. Telson ovate, posteriorly sparsely setose, apically broadly rounded.

Basal antennular peduncle article subequal in length but broader than 2 distal
articles; flagellum of 3 articles, second article longest, terminal article bearing 2
aesthetascs. Antennal peduncle article 2 grooved, article 5 longer than 4; flagellum
of 4 articles, together shorter than peduncle article 5. Mandibular palp with basal

626 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

E

_ Fig. 1. Apanthura zeewykae: A, Non-ovigerous °, dorsal view; B, Antenna; C, Antennule; D,
Maxilla; E, Maxilliped; F, Mandible.

article two-thirds length of article 2; terminal article one-third length of article 2,
bearing 5 distal spines; incisor of 3 cusps; lamina dentata of 6 serrations; molar
low, rounded. Maxilliped article 5 set obliquely on 4, with 4 distal setae; short
tapering endite on inner surface reaching distal margin of article 4 tipped with
single seta.

Pereopod 1 carpus with poorly developed broad tooth distally; propodus 1.5 x
as long as broad, palm with obscure tooth at midpoint, sparsely setose; unguis

VOLUME 95, NUMBER 3 627

Fig. 2. Apanthura zeewykae: A, Pereopod 1; B, Pereopod 2; C, Pereopod 7; D, Pleopod 1; E,
Telson; F, Uropodal exopod; G, Uropodal endopod and protopod.

half length of dactylus, with small accessory spine. Pereopod 2, carpus bearing
fringed scales on free posterior margin; propodus gently curved, with strong
posterodistal sensory spine, posterior margin bearing fringed scales; unguis less
than half length of dactylus. Pereopods 4—7, merus with strong anterior lobe but
no posterior lobe; carpus slightly lobed posteriorly; propodus 3 x as long as broad,
linear; unguis one-third length of dactylus, accessory spine one-fourth length of

628 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

unguis. Pleopod 1 operculiform, endopod slightly less than half width and five-
sixths length of exopod. Uropodal exopod 1.6 as long as broad, distally acute,
outer margin distally sinuate, setose, reaching almost to midlength of endopod;
latter longer than wide, distally rounded, setose, reaching beyond telsonic apex.

Material.—Holotype, AM-P32392, non-ovig. 2, TL 3.7 mm. Paratypes, AM-
P32393, 5 juveniles; NMV-J1707, non-ovig. 2, TL 3.5 mm, 5 juveniles; USNM
189062, non-ovig. 2, TL 3.3 mm, 5 juveniles; Rat Island, reef crest coral rubble,
1.3 m, 24 August 1981.

Remarks.—As noted in the past (e.g. Kensley 1979:817), the species of Apan-
thura tend to be separated on subtle morphological features. Shape of the uro-
podal rami, telson, degree of fusion of the pleonites, and maxillipedal proportions
seem to be the most useful characters in species separation. The present species
is similar to A. mana Kensley from Fiji, especially in pereopodal structure and
proportions, and in fusion of pleonites 1-5. The most distinctive feature of A.
zeewykae 1s the relatively strong rostrum, not seen in any previously described
species. Further support for the formation of a new species lies in the shape of
the uropodal rami, which are less elongate than in A. mana, and in the proportions
of the segments of the maxilliped and mandibular palp.

Etymology.—This species is named for the Dutch wooden gunship ‘‘Zeewyk,’’
wrecked on the Houtman Abrolhos Islands in 1727.

Eisothistos bataviae, new species
Figs. 3, 4

Description.—Integument posteriorly indurate. Dorsolateral eyes consisting of
8—9 ommatidia. Pleonites 1—5 short, similar; pleonite 6 twice length of 5. Telson
widening and broadly rounded posteriorly, with broad hyaline margin, lacking
dorsal spination, anterior half of lateral margin with row of short spines, posterior
margin becoming strongly serrate.

Antennule of 6 undifferentiated articles, terminal article bearing 2 aesthetascs.
Antenna with (?)6-articulated flagellum, not well demarked from peduncle. Man-
dible lacking palp, consisting of broadly rounded incisor of 3 faintly indicated
sclerotized cusps; proximal ridge present near base of appendage. Maxilla with
2 broad and 5 narrow terminal spines. Maxilliped slender-elongate, with 6 setae
on short terminal segment, proximal segmentation obscure.

Pereopod | with basis, ischium, and merus each bearing 3 very elongate setae;
carpus triangular; propodus not expanded, length about twice width; unguis one-
third length of rest of dactylus, slightly hooked. Pereopod 2, basis with 4 very
long setae; ischium with single elongate seta; carpus triangular, sparsely setose;
propodus bearing 6 short spines on posterior margin; unguis distally bearing mi-
nute spinules, about one-fourth length of dactylus. Pereopods 4—7, basis and
ischium bearing very elongate setae; carpus with anterior margin somewhat short-
er than posterior, latter with posterodistal spine and fringed scales; propodus
elongate-rectangular, with strong posterodistal sensory spine, posterior margin
bearing fringed scales; unguis about one-third length of dactylus. Rami of pleopod
1 fused, with short slit in distal margin, bearing 5 plumose setae on mesial margin,
12 plumose setae on distal margin. Uropodal exopod with anterior margin round-
ed, setose; strong elongate lobe at midlength of outer margin, armed with 4 short

VOLUME 95, NUMBER 3 629

Fig. 3. Eisothistos bataviae: A, Antennule; B, Antenna; C, Mandible; D, Maxilla; E, Maxilliped;
F, Uropodal endopod and protopod; G, Telson; H, Pleopod 1; I, Uropodal exopod.

spines on dorsal surface; posterior half of outer margin strongly toothed; uropodal
endopod wider than long, margins very strongly toothed; protopod with 8 elongate
plumose setae on dorsal surface, outer distal angle serrate.
Material.—Holotype, AM-P32395, non-ovig. 2, TL 3.0 mm; Paratypes, AM-
P32396, non-ovig. 9, TL 2.0 mm; NMV-J1718, 2 non-ovig. 2°, TL 2.0, 2.3 mm;

630 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Eisothistos bataviae: A, Pereopod 1; B, Pereopod 2; C, Pereopod 5.

USNM 189063, 2 non-ovig. 2, TL 2.5, 2.7 mm; Rat Island, reef crest coral rubble,
1.3 m, 24 August 1981.

Remarks.—Eisothistos bataviae shares with other species in the genus serrate
margins on the uropod and telson, and the well-developed spine on the uropodal
exopod. In these characters the genus is similar to Stellanthura Wagele, but
differs in having fused rami on pleopod 1.

Eisothistos bataviae is most similar to E. anomala (Kensley) (originally placed
in Heteranthura), but differs in the number of setae on pleopod 1, the number of
articles on antennae, and spination of the propodi. Eisothistos bataviae differs
from E. crateris Kensley, E. macrurus Wagele, and E. pumilis Wagele, in absence
of a middorsal spine row on the telson, and in possessing a serrate mesial margin
on the uropodal endopod. A dorsally smooth telson is found in E. maledivensis
Wagele, and in the only other Australian species, E. vermiformis Haswell. The
present species differs from these in possessing a more setose pleopod |. It differs
from all other species known in possessing long setae on articles 2 and 3 of the
pereopods.

There were numerous small serpulid worms in the samples from which the
Eisothistos specimens were taken. Wagele (1981) records these worms as the
prey of Eisothistos.

Etymology.—tThe specific epithet is derived from the name of a wooden vessel
of the Dutch East India Company, “‘Batavia,’’ wrecked on the Houtman Abrolhos
Islands in 1629.

VOLUME 95, NUMBER 3 631

ion

Fig. 5. Heptanthura sp.: A, Uropodal endopod; B, Uropodal exopod; C, Mandible.

A

Heptanthura sp.
Fig. 5

Material.—1 non-ovig. 2, TL 3.9 mm, AM-P32397; Rat Island, coral rubble
from reef crest, 1.3 m, 24 August 1981.

Remarks.—A few differences between the present specimen and H. novae-
zealandiae Kensley, 1978, could be detected. The mandibular palp proportions
differ slightly, the second article being longer in the New Zealand material, while
the terminal article carries more spines in the present specimen. The propodi of
the three posterior pereopods carry a few more spines distally, and the uropodal
endopod is proximally broader in the Abrolhos specimen. Of the body segments,
only pereonite 5 is slightly longer than 4 in the present specimen. The paucity of
material does not allow thorough comparison of this specimen with the type-
species of the genus, but given the geographical and ecological differences, it
seems unlikely that the two are conspecific.

Mesanthura protei Kensley
Fig. 6

Mesanthura protei Kensley, 1980:30, figs. 22, 23.

Material.—2 non-ovig. 2, TL 7.0 mm, 1 36, TL 4.0 mm, AM-P32398; Rat
Island, coral rubble from reef crest, 1.3 m, 24 August 1981.

Previous records.—Inhambane, Mozambique, 37 m; Nosi Bé, Madagascar, in-
tertidal to 1.5 m.

Remarks.—The appendages of both the male and female of the present material

632 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

\"

C

Fig.6. Mesanthura protei: A, Non-ovigerous 2 , dorsal view; B, Pereopod 2; C, Uropodal exopod.

agree completely with the original description. The dorsal pigment pattern of the
female, with its more or less complete rings on pereonites 1—6 and on the pleon,
and the solid bar across the posterior half of pereonite 7, is closest to that figured
by Kensley (1980) in his Fig. 22b.

Mesanthura protei differs from M. albinotata Thomson and M. bipunctata
Thomson from Rottnest Island in its pigment pattern and the shape of the uro-
pods.

This record extends the range of the species across the width of the Indian
Ocean, an unusually broad distribution for any shallow water anthurid.

The second pereopod and the uropodal exopod are figured here, as they were
not shown in the original description.

VOLUME 95, NUMBER 3 633

F

Fig. 7. Panathura haddae: A, Antenna; B, Antennule; C, Mandible; D, Maxilla; E, Maxilliped;
F, Uropodal endopod and protopod; G, Telson; H, Uropodal exopod; I, Pleopod 1.

Panathura ardea (Poore and Kensley)

Coralanthura ardea Poore and Kensley, 1981:507, figs. 3, 4.

Material.—1 ovig. 2, TL 3.2 mm, 2 non-ovig. 2, TL 3.6, 3.2 mm, AM-P32394;

west side of sandy shoal, Rat Island, from coral rubble and algal washings, 3—5
m, 25 August 1981.

Previous records.—Heron Island, southern Great Barrier Reef, Queensland,
Australia, intertidal.

Remarks.—This species was originally placed by us in a new genus of which

634 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 8. Panathura haddae: A, Pereopod 1; B, Pereopod 2; C, Pereopod 7.

the type is Coralanthura endeavourae Poore and Kensley. In this we were clearly
in error. Although the two genera are similar, Panathura contrasts with Cor-
alanthura in possession of more articles on antennae | and 2, a shorter mandibular
palp, a much narrower maxillipedal palp of which the terminal article is not
minute, absence of an anterior spine on the merus of pereopods and an almost
operculiform pleopod | with shortened tapering endopod. Most species of Pa-
nathura, though not the type-species P. serricauda Barnard, have a toothed palm
on pereopod | such as is seen in P. ardea. The only difference detected between
the present material and the original description lies in the mandible. The lamina
dentata has a finely serrulate part not seen in the Heron Island specimens, while
the terminal palp article bears one more spine. The flagellum of antenna 1 (both
in this and in the Queensland material) has four articles, not three as in the original
description. Each, except the first, bears a single aesthetasc.

Panathura haddae, new species
Figs. 7, 8

Description.—Integument relatively indurate in posterior half of body. Body
proportions: C = 1 >2 =3 <4=5 > 6 >7. Head with triangular rostrum barely
overreaching anterolateral lobes; dorsolateral eyes small. Pereonite 7 laterally
overlapping pleon. Pleonites free, 1-4 short, 5 twice length of 4, 6 subequal to
anterior pleonites together, with posterior margin middorsally broadly triangular.
Telson posteriorly broadly rounded, bearing several setae.

Antennule with basal peduncular article equal in length but wider than 2 distal
articles; flagellum of 3 articles. Antenna with second peduncular article subequal

VOLUME 95, NUMBER 3 635

in length to articles 3 and 4 together; article 5 longer than 4; flagellum of 3 articles.
Mandibular palp with article 2 longer than 1 or 3, with strong distal spine, terminal
article bearing 4 serrate spines; incisor of 3 cusps, lamina dentata consisting of
distal coarsely serrate and proximal finely serrate parts; molar reduced, low,
rounded. Maxilliped with narrowly triangular endite reaching penultimate palp
article, bearing 3 setae; antepenultimate article with strong distal serrate spine.

Pereopod | subchelate; carpus with posterodistal angle rounded, bearing trans-
parent flange; propodus expanded, palm with delicate transparent flange consist-
ing of 3 rounded lobes; unguis about half length of rest of dactylus. Pereopods 2
and 3 subchelate; carpus with posterodistal rounded flange; propodus not as ex-
panded as in pereopod 1, with transparent palmar flange having 2 low triangular
teeth; unguis half length of rest of dactylus. Pereopods 4—7, carpus with anterior
margin slightly shorter than posterior margin, latter with single spine and seta;
propodus with sensory spine at midlength and distal angle of posterior margin,
and 2 serrate spines at anterodistal angle; unguis only slightly less than dactylar
length. Pleopod 1 exopod operculiform, broadly ovate, with about 5S plumose
setae on distal margin; endopod just reaching distal half of, and about one-fourth
width of exopod. Uropodal exopod roughly ovate, with hyaline margin, anteriorly
and posteriorly rounded, with low setose lobe on outer margin; endopod broader
than long, with several elongate setae distally; protopod with 4 elongate plumose
setae on outer margin.

Material.—Holotype, AM-P32399, ovig. 2 , TL 2.4mm; Paratypes, AM-P32400,
3 juveniles; NMV-J1719, non-ovig. 2, TL 2.1 mm, 1 juvenile; USNM 189064,
non-ovig. 2, TL 2.8 mm, | juvenile; Rat Island, reef crest rubble, 1.3 mm, 24
August 1981.

Remarks.—Panathura haddae may be distinguished from the four species re-
corded from the Indo-Pacific region by the following features: the telson is more
broadly rounded posteriorly and is not serrate as in P. amstelodami Kensley,
while the uropodal exopod and endopod are broader; the exopod of pleopod 1 is
longer and narrower, and the endopod relatively longer in P. macronesia Kens-
ley, recorded from Mauritius and Madagascar; the western Indian Ocean species
also has one article less in the antennular and antennal flagella. Panathura ser-
ricauda Barnard possesses a serrate telson, while the uropodal endopod is longer
than wide. Panathura collaris Kensley, from the Cook and Fiji islands, has a
more elongate endopod of pleopod 1, the uropodal exopod is longer than wide,
the palm of pereopod 2 has more serrations, and the antennular and antennal
flagella possess fewer articles. The species may be separated from P. ardea
(Poore and Kensley) with which it co-occurs by the much broader uropodal rami
and telson.

Etymology.—The species is named for the German barque ‘‘Hadda’’ which
was lost on the Houtman Abrolhas Islands in 1877.

Acknowledgments

We thank Patricia Hutchings for collecting the material on which this paper is
based, and T. E. Bowman, Smithsonian Institution, for reading and commenting
on the manuscript.

636 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Literature Cited

Kensley, B. 1978. Five new genera of anthurid isopod crustaceans.—Proceedings of the Biological
Society of Washington 91:775—792.

—. 1979. New species of anthurideans from the Cook and Fiji Islands (Crustacea: Isopoda:
Anthuridea).—Proceedings of the Biological Society of Washington 92:814—836.

——. 1980. Anthuridean isopod crustaceans from the International Indian Ocean Expedition,
1960-1965, in the Smithsonian collections.—Smithsonian Contributions to Zoology 304: 1-37.

Poore, G. C. B., and B. Kensley. 1981. Coralanthura and Sauranthura, two new genera of an-
thurideans from northeastern Australia (Crustacea: Isopoda: Anthuridae).—Proceedings of the
Biological Society of Washington 94:503—513.

Thomson, J. M. 1946. New Crustacea from the Swan River estuary.—Journal of the Royal Society
of Western Australia 30:35-73.

—. 1951. The Fauna of Rottnest Island X. Anthuridae.—Journal of the Royal Society of West-
ern Australia 35:1-18.

Wagele, J. W. 1981. Zur Phylogenie der Anthuridea (Crustacea, Isopoda) mit Beitragen zur Le-
bensweise, Morphologie, Anatomie und Taxonomie.—Zoologica 132:1—127.

(BK) Department of Invertebrate Zoology, National Museum of Natural His-
tory, Smithsonian Institution, Washington, D.C. 20560; (GCBP) Department of
Crustacea, National Museum of Victoria, Victoria Crescent, Abbotsford, Vic-
toria, Australia.

PROC. BIOL. SOC. WASH.
95(3), 1982, pp. 637-638

COMMENTS ON THE NOMENCLATURAL STATUS OF THE
FAMILIES CAUCASELLIDAE AND FAVUSELLIDAE
(FORAMINIFERIDA)

Richard W. Huddleston

Abstract.—The taxonomic nomenclature of the foraminiferal families Cauca-
sellidae and Favusellidae is examined. The nomenclatural complexities and status
of the genera Caucasella Longoria, 1974, Polskanella Fuchs, 1973, and Globu-
ligerina Bignot and Guyader, 1971 are reviewed in relation to the rules of the
International Code of Zoological Nomenclature.

Bignot and Guyader (1971) established Globuligerina as a new subgenus of
Globigerina d’Orbigny, 1826, designating Globigerina oxfordiana Grigelis, 1958,
emend. Bignot and Guyader, 1967, emend. Bignot and Guyader, 1971 as the type-
species. Fuchs (1973) considered Globigerina oxfordiana Grigelis, 1958 sensu
Bignot and Guyader 1967 and 1971 to represent a distinct form from Globigerina
oxfordiana Grigelis, 1958. Fuchs (1973) designated Globigerina oxfordiana Gri-
gelis, 1958 as the type-species of Polskanella Fuchs, 1973, and selected Globu-
ligerina frequens Fuchs, 1973 as a ‘‘replacement’’ type-species for Globuligerina.
Grigelis and Gorbatchik (1980:184) criticized Fuchs (1973) and commented, ‘‘In
‘as much as the type of a taxon indicated in accordance with the ICZN [Inter-
national Code of Zoological Nomenclature] regulations is not subject to replace-
ment, Fuchs’ choice of a new type species for the genus Globuligerina is illegal,
and Polskanella as a genus based on an already used type species should be
assigned to younger objective synonyms of Globuligerina (ICZN, 1966, p. 35
article 61b).”’

The significant fact that has been overlooked is that Bignot and Guyader (1971)
specifically designated Globigerina oxfordiana Grigelis, 1958 emend. Bignot and
Guyader 1967, emend. 1971, and not Globigerina oxfordiana Grigelis, 1958. Big-
not and Guyader (1971) are considered to have established a new nominal species,
with the same specific name as the misidentified species, in the new nominal
genus, Globuligerina (ICZN, Article 70(b)i). Therefore, the type-species of Glob-
uligerina is to be cited as Globuligerina oxfordiana Bignot and Guyader, 1971
(ICZN, Article 70b).

The specific name Globuligerina oxfordiana Bignot and Guyader, 1971 cannot
be retained for the species used as the type-species of Globuligerina (ICZN,
Article 49). I, herein, propose the new species Globuligerina conflicta for the
Specimens currently identified as Globuligerina oxfordiana Bignot and Guyader
(=Globigerina oxfordiana Grigelis sensu Bignot and Guyader not G. oxfordiana
Grigelis). Whether or not Globigerina oxfordiana Grigelis or Globuligerina con-
flicta is the type-species of Globuligerina is a matter that can be resolved only by
the International Commission on Zoological Nomenclature (ICZN, Article 70a).

Polskanella Fuchs complicates the problem. This genus may be a junior ob-
jective synonym of Globuligerina (ICZN, Article 61b) if the Commission rules
Globigerina oxfordiana Grigelis as the type-species of Globuligerina. However,
if the Commission rules Globuligerina conflicta as the type-species of Globulig-
erina, then Polskanella would be firmly established.

638 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fuchs’ (1973) designation of Globuligerina frequens Fuchs as the new type-
species of Globuligerina has no standing since G. frequens is distinct from Glo-
bigerina oxfordiana Grigelis sensu Bignot and Guyader.

Longoria (1974) established two families, Caucasellidae (type-genus Caucasella
Longoria, 1974) and Favusellidae (type-genus Favusella Michael, 1972). Longoria
(1974) designated Globigerina hauterivica Subbotina, as the type-species of Cau-
casella and included 6 other species within this genus, including Globigerina
oxfordiana. The generic descriptions and included species of Caucasella and
Globuligerina are nearly identical and these genera were considered congeneric
by Grigelis and Gorbatchik (1980). Grigelis and Gorbatchik (1980) placed Glob-
uligerina within the Favusellidae, rendering Caucasellidae a junior synonym of
Favusellidae.

The name Caucasella previously was used by Moisseev (1934) as a genus of
Jurassic brachiopod. Caucasella Longoria is a junior homonym which renders
the family Caucasellidae Longoria invalid (ICZN, Article 39). A new name (no-
men novum) for Caucasella Longoria is not necessary due to its status as a junior
synonym, and is replaced with the oldest available synonym (ICZN, Article 60a).
The oldest available synonym to replace Caucasella Longoria would be Polska-
nella if Polskanella and Globuligerina are held distinct. However, if Polskanella
and Globuligerina are viewed congeneric, then Globuligerina would be the re-
placement name for Caucasella.

Acknowledgments

I thank K. L. Finger and D. Haman, Chevron Oil Field Research Company
for review of this manuscript and providing valuable comments.

Literature Cited

Bignot, G., and J. Guyader. 1967. Découverte de foraminiferes planctoniques dans |’Oxfordien du

Havre (Seine-Maritime).—Revue de Micropaléontologie 9(2):104—110.

, and 1971. Observations nouvelles sur Globigerina oxfordiana Grigelis.—Proceed-

ings of the II Planktonic Conference (Rome, 1970):79-83.

Fuchs, W. 1973. Ein Beitrag zur Kenntnis der Jura-Globigerinen und verwandten Formen an Hand
polnischen Materials der Collovien und Oxfordien.—Verhandlungen der Geologischen Bun-
desanstalt 3:445—487.

Grigelis, A. A. 1958. Globigerina oxfordiana sp. nov. a discovery of Globigerina in the upper

Jurassic deposits of Lithuania.—Transactions of the Academy of Sciences U.S.S.R. 219(5):

1203-1205.

, and T. Gorbatchik. 1980. Morphology and taxonomy of Jurassic and Early Cretaceous

representatives of the superfamily Globigerinacea (Favusellidae)—Journal of Foraminiferal

Research 10(3): 180-190.

Longoria, J. F. 1974. Stratigraphic, morphologic and taxonomic studies of Aptian planktonic fo-
raminifera.—Revista Espanola Micropaleontologica Numero Extraordinario, 107 pp.

Michael, F. Y. 1972. Planktonic Foraminifera from the Comanchean Series (Cretaceous) of Texas.—
Journal of Foraminiferal Research 2(4):200-—220.

Moisseev, A. S. 1934. Brakhiopody Iurskikh otlozhenii Krym i Kavkasa.—Vsesoiuznoe Geologo-
Razvedochnoe Otdelenie Trudy v. 203, 213 pp.

Orbigny, A. D.d’ 1826. Tableau méthodique de la classe des Cephalopodes.—Annales des Sciences
Naturelles, Paris, ser. 1, 7:245-314.

Chevron Oil Field Research Company, P.O. Box 446 La Habra, California
90631. (Present address: Scientific Research Systems 11044 McGirk, El Monte,
California 91731.)

BB

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CONTENTS

Systematic resolution of the genera of the Crinia complex (Amphibia: Anura: Myobatrachidae) —
W. Ronald Heyer, Charles H. Daugherty, and Linda R. Maxson

Mexican freshwater silversides (Pisces: Atherinidae) of the genus Archomenidia, with the de-

scription of a new species Barry Chernoff and Robert Rush Miller

A new species of skate from the western Indian Ocean, with comments on the status of Raja
(Okamejei) (Elasmobranchii: Rajiformes) John D. McEachran and Janice D. Fechhelm
Relationships of the African killifish genus Foerschichthys (Teleostei: Cyprinodontiformes:
Aplocheilidae) Lynne R. Parenti
A new xystodesmid milliped genus and three new species from the eastern Blue Ridge Moun-
tains of North Carolina (Polydesmida) Rowland M. Shelley
Naushonia panamensis, new species (Decapoda: Thalassinidea: Laomediidae) from the Pacific
coast of Panama, with notes on the genus Joel W. Martin and Lawrence G. Abele
Paralvinella grasslei, new genus, new species of Alvinellinae (Polychaeta: Ampharetidae) from
the Galapagos Rift geothermal vents Daniel Desbruyéres and Lucien rae
A new genus of bomolochid copepods from Indo-West Pacific nemipterid fishes

nation in the ophiuroid family Ophiacanthidae (Echinodermata: Ophiuroidea) from off

Oregon, U.S.A. Michael A. Kyte

Mesonerilla prospera, a new po from marine caves in Bermuda
Wolfgang Sterrer and Thomas M. Iliffe

Two new species of Ceratonereis (Polychaeta: Nereididae) from estuarine areas of New

South Wales, Australia P. A. Hutchings and C. J. Glasby
Anopsilana crenata, a new troglobitic cirolanid isopod from Grand Cayman Island, Carib-
bean Sea Thomas E. Bowman and Richard Franz
Uncispionidae, a new polychaete family (Annelida) Karen D. Green

Four new species of stomatopod crustaceans from the Philippines
Renato G. Garcia and Raymond B. Manning
A new crayfish of the genus Procambarus from southwestern Arkansas
Horton H. Hobbs, Jr., and Henry W. Robison
A new Uloborus Latreille species from Argentina (Arachnida: Araneae: Uloboridae)
Brent D. Opell
A new genus and species of poison-dart frog (Amphibia: Dendrobatidae) from the Andes

of northern Colombia John D. Lynch and Pedro M. Ruiz-Carranza —

A note on the genitalia of Potamothrix hammoniensis (Oligochaeta: Tubificidae) H.R. Baker
Freshwater triclads (Turbellaria) of North America. XIV. Polycelis monticola, new species,
from the Sierra Nevada range in California Roman Kenk and Anne M. Hampton
Redescription of the major spines of Polydora ligni Webster (Polychaeta: Spionidae)
Jerry D. Kudenov
Astreptosyllis acrassiseta, a new genus and species of the subfamily Eusyllinae (Polychaeta:
Syllidae) from Australia Jerry D. Kudenov and John H. Dorsey
Pinnixa costericana, a new species of crab from Central America (Brachyura: Pinnotheridae)
| Mary K. Wicksten
Review of some little-known species of syllids (Annelida: Polychaeta) described from the Gulf of
Mexico and Caribbean by Hermann Augener in 1924 Joan M. Uebelacker
The status of two Chilean frogs of the genus Eupsophus (Anura: Leptodactylidae)
J. R. Formas and M. Inés Vera
Morphology and development of planktonic Lolliguncula brevis (Cephalopoda: Myopsida)
Michael Vecchione
Redescription of Exoediceros fossor (Stimpson, 1856) an Australian marine fossorial ree
the type-genus of the new family Exoedicerotidae
J. Laurens Barnard and Margaret M. Drummond

Courtship display in a Bornean frog Keith A. Harding
Anthurids from the Houtman Abrolhos Islands, Western Australia (Crustacea: Isopoda:
Anthuridae) Brian Kensley and Gary C. B. Poore

Comments on the nomenclatural status of the families Caucasellidae and Favusellidae
(Foraminiferida) _ Richard W. Huddleston

Roger F. Cressey
Ophiacantha abyssa, new species, and Ophiophthalmus displasia (Clark), a suggested new combi- |

423
428

440

451

458
478
484

495
505

509,

oh eh

‘500.

530
537
545
554

557
563

567
571
575
579
583
594
602
610
621
625

637

| i he as r pose a. i ! } (ISSN 0006-324X)

O

/ASHINGTON

(20 December 1982 Number 4

4

THE BIOLOGICAL SOCIETY OF WASHINGTON

1982-1983
Officers
President: Paul J. Spangler | mi Secretary: Catherine J. Kerby
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PROCEEDINGS
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PS et eee ee, Se a ee See

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PROC. BIOL. SOC. WASH.
95(4), 1982, pp. 639-641

PTERYNOTUS XENOS, A NEW SPECIES OF MURICID
FROM OFF NORTHERN JAMAICA
(MOLLUSCA: GASTROPODA)

M. G. Harasewych

Abstract.—A new muricid, Pterynotus xenos, is described from 60 meters off
Discovery Bay, Jamaica. The habitat of this new species is discussed, as are its
relationships with closely related Recent and fossil species.

During the preparation of a review of the Pterynotus of the western Atlantic
(Harasewych and Jensen 1979), the authors were aware of a report of Pterynotus
phaneus Dall, 1889 [as Pterynotus tristichus Dall, 1889] occurring in 70 meters
off Discovery Bay, Jamaica (Humfrey 1975:136, pl. 22, fig. 35). As this report
was not consistent with previously recognized geographic and bathymetric ranges
of this species, and since the illustration cast doubt on the identification, we felt
it best not to include the record under P. phaneus.

Since that time, additional specimens from the same locality were brought to
_ my attention by Dr. Emily H. Vokes. Examination of this material has shown
that the specimens are not referable to Pterynotus phaneus, but represent a new
species, which is described here.

Family Muricidae
Subfamily Muricinae
Genus Pterynotus Swainson, 1833

Pterynotus (Pterynotus) xenos, new species
Figs. 1-3

Description.—Shell small (to 7 mm), fusiform, moderately heavy for size; spire
angle 37—42°; protoconch of one and one-quarter whorls, low, pitted, ending in a
distinct, thin varix; teleoconch with 4 convex whorls; 3 thin, broad varices per
whorl; first postnuclear whorl smooth between varices; second and third post-
nuclear whorls with 2 to 3 intervarical nodes of equal size; fourth postnuclear
whorl with 2 intervarical nodes, which may be unequal in size; spiral sculpture
of 5 to 7 major cords with numerous fine spiral threads throughout; laminae on
ventral surfaces of varices very fine; aperture oval; inner lip smooth, attached
posteriorly; outer lip smooth or with 4 denticles; siphonal canal of moderate
length, open, straight, tip slightly recurved dorsally; shell color pale salmon;
operculum, periostracum, and soft parts unknown.

Type-specimens.—Holotype, USNM 703309, length 6.30 mm; Paratype 1,
USNM 784590, length 6.15 mm; Paratype 2, USNM 784590, length 5.08 mm.

Type-locality.—Off Discovery Bay, Jamaica, 60 meters.

Range.—At present known only from the type-locality.

Material examined.—The 3 type-specimens.

Ecology.—Collected from corals on reef wall. Humfrey (1975) reported col-
lecting 2 specimens from a blade of Agaricia coral at the type-locality.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-3. Pterynotus (Pterynotus) xenos: 1, Holotype, USNM 703309, (10.0x); 2, Paratype 1,
USNM 784590, (10.0x); 3, protoconch of paratype 2, USNM 784590. Scale bar equals 250 um.

VOLUME 95, NUMBER 4 641

Etymology.—From xenos Gr.—stranger. The name is proposed as a noun.

Remarks.—Although Pterynotus xenos superficially resembles, and has been
confused with P. phaneus, it may readily be distinguished from the latter by its
smaller, thicker shell, its thickened varices which are buttressed on the apertural
side, its prominent spiral sculpture as well as by its salmon color. These two
species are members of lineages that have been distinct since the Middle Miocene.
Pterynotus phaneus traces its origin to the Helvetian P. delaunayi (Tour-
nouer,1875). All known Recent species in this lineage are members of upper
continental slope communities. Pterynotus xenos is most similar to P. venustus
(Bellardi, 1872) from the Miocene of Italy, both in appearance and habitat. As
more material becomes available, the nearest living relative of P. xenos will likely
prove to be P. tripterus (Born, 1778) from the Indo-Pacific.

Chicoreus cosmani Abbott and Finlay, 1979, another recently described mur-
icid from the reefs off Ocho Rios, Jamaica, is also more similar to Chicoreus
dujardini (Tournouer, 1875) from the Helvetian of France than to any Recent
species. In the light of recent discoveries of relict pockets throughout the Carib-
bean (Petuch 1981a, b), the offshore reef fauna of northern Jamaica would seem
to merit further investigation because it seems to substantiate this trend.

Acknowledgments

I would like to thank Dr. Emily H. Vokes of Tulane University for making this
material available for study. I also thank Dr. Vokes as well as Dr. Edward J.
Petuch of Continental Shelf Associates, Jupiter, Florida, for critical review of the
manuscript.

Literature Cited

Harasewych, M. G., and R. H. Jensen. 1979. Review of the subgenus Prerynotus (Gastropoda:
Muricidae) in the western Atlantic.—Nemouria 22: 1-16.

Humfrey, M. 1975. Sea shells of the West Indies. —Taplinger Publishing Co., New York. 351 pp.
Petuch, E. J. 198la. A relict Neogene caenogastropod fauna from northern South America.—
Malacologia 20(2):307—347.

1981b. A volutid species radiation from northern Honduras with notes on the Honduran
Caloosahatchian secondary relict pocket.—Proceedings of the Biological Society of Washing-
ton 94(4):1110—-1130. }

College of Marine Studies, University of Delaware, Newark, Delaware 19711.

PROC. BIOL. SOC. WASH.
95(4), 1982, pp. 642-646

ISTIGOBIUS HOESEI, A NEW GOBIID FISH FROM
AUSTRALIA (PERCIFORMES: GOBITDAE)

Edward O. Murdy and John D. McEachran

Abstract.—A new species of Istigobius is described based on 18 specimens
from waters near Sydney, Australia, the only known locality for this species. It
differs from all other nominal species of /stigobius in its distinctive coloration.
The new species is most similar to J. campbelli, a species from Japan, China,
and Taiwan.

The gobiid taxon Istigobius Whitley was elevated to generic status by Hoese
and Winterbottom (1979). Prior to this it was widely accepted as, and commonly
found in the literature as, Acentrogobius. The genus is distributed from the Red
Sea to the Marquesas Islands in the Central Pacific. Species of [stigobius are
sand-dwelling, many being found associated with coral reefs. During the course
of a generic revision, we encountered a new species that differs strikingly in
coloration from all nominal forms. At present this species has only been collected
from waters around Sydney, Australia, and is herein described from 18 speci-
mens.

Methods

Except as indicated, counts and measurements were taken from the left side
and follow methods given by Hubbs and Lagler (1958). Head length is taken to
the upper attachment of the opercular membrane. In most gobiids the first ray of
the second dorsal and anal fins is simple and is included in ray counts. The last
‘ray of each of these fins is branched at the base and is counted as a single element.
The longitudinal scale count is taken from the upper attachment of the opercular
membrane to the end of the hypural. Transverse scale counts are taken from the
anal origin upward and forward to the first dorsal base (TRF) and from the anal
origin upward and backward to the second dorsal fin base (TRB). Since the
anterior scales are often crowded and irregularly placed, the TRB count generally
shows less variation (Hoese and Steene 1978). Predorsal scales extend from just
anterior to the first dorsal spine to just posterior to the interorbital region and are
counted in a straight-line manner. Postdorsal scales are counted from the end of
the second dorsal fin to the upper base of the caudal fin. The vertebral count
includes the urostyle. Pectoral fin rays were counted on both sides and tabulated
as separate counts.

We follow the methods of Birdsong (1975) in describing the relationship of the
spinous dorsal fin pterygiophores with the underlying vertebrae.

All measurements less that 20 mm were made to the nearest 0.01 mm with an
ocular micrometer. Measurements greater than 20 mm were made to the nearest
0.1 mm with dial calipers. All fish lengths given are standard lengths (SL).

In the descriptive accounts the values for meristic characters are followed by
frequencies of counts in parentheses. The frequency that includes the value for
the holotype is italicized.

VOLUME 95, NUMBER 4 643

Fig. 1. Male paratype of Istigobius hoesei, AMS 1.17657-001, 59.5 mm SL. Photograph by E. O.
Murdy.

Istigobius hoesei, new species
Figs ig2

Holotype.—USNM 231865, male, 54.9 mm, 33°50’S 151°16’E, Sydney Harbour,
Australia, collected by R. Kuiter on 6 Sept. 1981 with rotenone at a depth of
5—7 m in sandy area near large rocks.

Paratypes.—AMS I. 17657-001, males (54.5 and 59.5), females (40.7, 49.5, 51.1,
51.2, 55.3, 56.1, 60.2, and 66.3), 34°00’S 151°14’E, Bare Island, La Perouse,
Botany Bay, New South Wales, R. Kuiter, 15 Dec. 1973; TCWC 3215-1, female
(38.6), same data as holotype; BPBM 27826, female (48.6), same data as holotype.

Diagnosis.—A species of moderate size (38.6—-66.3 mm SL) normally having
second dorsal of I, 10 and anal of I, 9 with pectoral rays usually 17-18. Pelvic
fins completely united, I, 5. Nape, breast and pectoral base covered with cycloid
scales. Body with 28—33 scale rows. TRB 6.5—8.5. Body irregularly covered with
dark spots and faint longitudinal lines. Lateral portions of jaws, especially upper
jaws, blackened. Two spots on pectoral base, uppermost one spreading out onto
fin rays. First spines and membranous portions of dorsal fins spotted. Males with
a flattened, tapering genital papilla, females possessing more bulbous type.

Description.—Dorsal rays VI-I, 10 (/7), VI-I, 11 (1); anal rays I, 8 (1), I, 9
(16), I, 10 (1); pectoral rays (right side) 14 (1), 16 (2), 17 (10), 18 (4), 19 (1);
pectoral rays (left side) 16 (2), 17 (9), 18 (5), 19 (1); segmented caudal rays 17
(/8); branched caudal rays 14 (7), 15 (9), 16 (2); lateral scale rows 28 (1), 29 (3),
30 (3), 31 (@), 32 (4), 33 (4); transverse scale rows (TRF) 7 (2), 7.5 (4), 8 (2), 8.5
(7), 9 (1), 9.5 (1); transverse scale rows (TRB) 6.5 (1), 7 (3), 7.5 (7), 8 (4), 8.5 (1);
circumpeduncular scales 12 (/6); postdorsal scales 5 (1), 5.5 (/), 6 (16); predorsal
scales 8 (1), 9 (2), 10 (9), 11 (3), 12 (1); gill rakers on outer face of first arch 1+4
(6), 1+5 (4). Vertebrae 10+ 16 (5); pterygiophore formula 3 (22110) (4 specimens),
3 (21210) (1). Morphometric data are given in Table 1.

644 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

SZ LEE EE

Wt LLL SE
ae Ze
2 ores

\ KN a ‘ \ se ANG iN 2

Fig. 2. Female paratype of Istigobius hoesei, AMS 1.17657-001, 49.5 mm SL. Drawing by Janice
D. Fechhelm.

Head slightly depressed, wider than deep. Body compressed. Mouth subter-
minal. Jaws extended posteriorly to a point under anterior portion of eye. Inter-
orbital very narrow. Anterior nostril a short tube above upper lip. Posterior nostril
a simple pore close to anterior margin of eye. Gill opening somewhat restricted,
ending at ventral midpoint of operculum. Gill rakers short, about as wide as long.
Dorsal and anal rays branched. Fourth spine of first dorsal fin longest. Pectoral
fin elongate with pointed tip reaching (in holotype) or almost reaching above anal
origin; all but 2 uppermost rays branched. Caudal fin rounded. Pelvics connected
by frenum, soft rays all branched; medial rays longest.

Two supraorbital canals (Fig. 3) each terminating medial to a nasal pore; a
medial anterior and medial posterior interorbital pore; one postocular pore behind
each eye; one infraorbital pore located posterio-ventrally from orbit; lateral canal
extending from posterior end of eye to middle of operculum, with one pore at
uppermost edge of preoperculum and a terminal pore above the middle of oper-
culum; a short tube with pores at each end above posterior portion of operculum;
three preopercular pores, lowest pore in horizontal line behind end of jaws.

No vomerine or palatine teeth. All jaw teeth conical and curved. Jaw teeth
tightly packed in 4-5 rows in both upper and lower jaws, outer row in each jaw

Table 1.—Proportional measurements of Istigebius hoesei in thousandths of standard length.

Paratypes
10 2 26
OCCOE__ _ _ __—_— oo
Character fe) Range Mean Range Mean

Standard length (mm) 54.9 38.6-66.3 Slo 7 2B 335) 54.5-59.5 57-02 3255
Proportions

Head length a3 .236-.270 .256 + .010 .250-.260 ape (QUT

Eye diameter .061 .055—.072 .064 + .005 .056—.061 059 + .004

Upper jaw length .088 .068—.087 .076 + .006 .079-.081 .080 + .001

Lower jaw length .076 .063—.080 .070 + .005 .073—.076 075 + .002

Body depth . 164 . 164—. 193 .174 + .009 .165—.165 .165 + .000

Head depth 182 .151-.180 .167 + .008" — .168—.180 174 + .009

Least depth caudal peduncle Zi .098—.122 .116 + .007 .118-.119 119 + .001

Pectoral length Vs .251—.290 PANG) ae KDI3) .275—.275 .275 + .000

Pelvic length .243 .215—.248 1235 =" OZ .250—.254 22h 008

VOLUME 95, NUMBER 4 645

Fig. 3. Pore and papillae patterns of Istigobius hoesei.

possessing the largest teeth. Upper jaw with 4 enlarged teeth on outer row, outer
row of lower jaw possessing 10 enlarged teeth with two most lateral curved
backward.

Body covered with 28-33 scale rows. Scales on nape, breast and pectoral bases
cycloid, all others ctenoid. Preoperculum and operculum naked. Isthmus scales
extend anteriorly to below midpoint of operculum.

Coloration of freshly preserved and photographed paratype (BPBM 27826)
showed head mostly greenish-brown with faint yellow-orange bands diagonally
on preoperculum and operculum. Lateral aspect of upper jaw black with some
blackened areas in preoperculum. Body with 5-6 greenish-brown longitudinal
lines on grayish-white background, reddish-orange spots and dashes dorsally.
Reddish-brown mid-lateral spots with 6-8 reddish-brown spots ventro-laterally.
Dorsal fins with red and white spots on first spines and on membranes. Orange
stripe basally on anal fin. Three horizontal bands on fleshy pectoral-fin bases.
Two brown spots on pectoral-fin bases, uppermost one spreading onto fin rays.
A metallic green glistening ventrally is sometimes retained in preserved speci-
mens. Caudal fin possessing some faint yellow streaks.

In preservation, head and body brownish. Gray, tan or white ventrally. Some
specimens with horizontal dark bars beneath eyes. Upper jaw black on posterio-
lateral aspect. Body mottled with irregular markings. Generally, 6-8 dark spots
discernible ventro-laterally. Two dark spots on pectoral-fin bases, uppermost one
spreading onto fin rays. Dark spots on first spines and membranes of dorsal fins.
Males with darkened pelvic and anal fins with elongation of soft dorsal and anal
fins, which is characteristic of most species of /stigobius examined.

Distribution.—Collected only in bays near Sydney, New South Wales, Austra-
lia.

Ecology.—The holotype and two paratypes were taken on sand near large rocks
at a depth of 5-7 m. Water temperature at the time of collection was 14°C.

Etymology.—The species is named for Douglass F. Hoese, for his contributions
towards a better understanding of gobioid systematics.

Relationships.—Istigobius hoesei is most similar to Istigobius campbelli (Jor-

646 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

dan and Snyder, 1901) which is found in Japan, China, and Taiwan. Istigobius
cambelli differs from I. hoesei in possessing a dark bar from the eye to the upper
pectoral-fin base, lacking blackened portions on the lateral aspects of upper jaw,
isthmus being scaled anteriorly to below distal portion of preoperculum, generally
deeper body, larger size, and more uniform pattern of spots. Field data included
with two specimens of I. campbelli collected in Taiwan by V. G. Springer (VGS
68-28) indicated these fishes were also collected near rocks at a depth of 5 m.
These Taiwanese specimens of [. campbelli are not as deep bodied as more
northerly conspecifics and are the most similar to J. hoesei, but the two species
are easily distinguishable by the characteristics given above. Istigobius hoesei is
easily distinguishable from all other [stigobius on color alone.

Acknowledgments

We acknowledge the curators and staffs responsible for the fish collections at
the Australian Museum, Sydney (AMS), Bernice P. Bishop Museum, Honolulu
(BPBM), and the National Museum of Natural History, Washington (USNM) for
the loan and exchange of study material and data. Rudie H. Kuiter collected all
type material. John E. Randall provided a color transparency of a fresh paratype.
Janice D. Fechhelm used her extensive talents to draw Figures 2 and 3. The
Lerner-Gray Fund for Marine Research of the American Museum of Natural
History partially supported this research.

Literature Cited

Birdsong, R. 1975. The osteology of Microgobius signatus Poey (Pisces: Gobiidae), with comments
on other gobiid fishes.—Bulletin of the Florida State Museum Biological Sciences 19(3):135-
187.

Hoese, D. F., and R. Steene. 1978. Amblyeleotris randalli, a new species of gobiid fish living in
association with alphaeid shrimps.—Records of the Western Australian Museum 6(4):379-389.

Hoese, D. F., and R. Winterbottom. 1979. A new species of Lioteres (Pisces, Gobiidae) from
KwaZulu, with a revised checklist of South African gobies and comments on the generic
relationships and endemism of western Indian Ocean gobioids.—Royal Ontario Museum, Life
Sciences Occasional Papers 31:1—13.

Hubbs, C. L., and K. F. Lagler. 1958. Fishes of the Great Lakes Region.—Bulletin of the Cranbook
Institute of Science, 26. 213 pp.

Department of Wildlife and Fisheries Sciences, Texas A&M University, Col-
lege Station, Texas 77843.

PROC. BIOL. SOC. WASH.
95(4), 1982, pp. 647-656

SUBSPECIES AND GEOGRAPHIC DISTRIBUTION OF
BLACK-MANTLE TAMARINS SAGUINUS
NIGRICOLLIS SPIX (PRIMATES: CALLITRICHIDAE)

Philip Hershkovitz

Abstract.—A new subspecies from Colombia of the black-mantle tamarin, Sa-
guinus nigricollis, is distinguished from S. n. nigricollis and S. n. graellsi by color
pattern. Geographic range of each of the three subspecies is revised and dis-
cussed, and differentiation of the taxa during early Recent—Late Pleistocene is
hypothesized on the basis of past climates.

Description of a new subspecies of black-mantle tamarin, herein named Sa-
guinus nigricollis hernandezi, entails some modifications of my 1977 account of
the species. As previously noted (Hershkovitz 1977:618, 626) no single character
or combination of characters other than color and color pattern consistently dis-
tinguishes any species or subspecies of tamarin of the Saguinus nigricollis group.
The distinctive color patterns of each of the three presently recognized races of
_ Saguinus nigricollis are given in the following key. For use of color and color
pattern terminology see Hershkovitz (1977:91-101).

Key to Subspecies of the Black-mantle Tamarin
(Fig. 1)

1. Nape and mantle blackish or dark brown (eumelanin) agouti, the mantle
more or less square behind and not extending beyond midback; remainder
of back, sides of body, thighs, legs, and proximal 5-16 cm of dorsal and
ventral surface of tail buffy (pheomelanin) agouti; remainder of tail black-
ish; neck, chest, and belly varying from dominantly buffy agouti to dom-
inantly blackish ....... Saguinus nigricollis graellsi Jiménez de la Espada.
2. Nape and mantle nearly or entirely uniformly blackish (eumelanin), the
mantle usually tapered behind and extending to lower back; sacral region
and dorsal surface of tail with proximal 3—5 cm marbled or striated reddish
(pheomelanin) and blackish; ventral surface of tail base, rump, and legs
dominantly or nearly. entirely reddish or mahagony, remainder of tail
blackish; neck, chest, and belly blackish, often mixed with reddish or
INE] NO IN, MS sk i ee Saguinus nigricollis nigricollis Spix.
3. Nape and mantle nearly uniformly blackish, the mantle tapered behind to
midback with the blackish continued, usually as a mid-dorsal band or
stripe, across lower back and proximal portion of tail; sides of lower back
mixed blackish and orange (pheomelanin), sides and ventral surface of
proximal 5-10 cm of tail orange agouti, remainder of tail blackish; neck
and chest dominantly orange agouti, belly mixed orange and blackish
3 2 die hth PN ie ed aria ere ea Saguinus nigricollis hernandezi, new subspecies.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

648

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VOLUME 95, NUMBER 4 649

Saguinus nigricollis hernandezi, new subspecies

Saguinus fuscicollis.—Izawa (not Spix), 1975:304, Fig. 5 (animal on tree trunk
eating resin); COLOMBIA: Caquetd (Rio Peneya); behavior (food, feeding).—
Izawa, 1976:384; COLOMBIA: Caquetd (Rio Peneya); behavior (social groups,
group sizes).

Saguinus nigricollis —Hernandez Camacho in Hershkovitz, 1977:1021; COLOM-
BIA: Caquetd (Curiplaya, Rio Caqueta).—Izawa, 1978:241, 274, fig. 4 (allo-
grooming), fig. 5 (autogrooming), fig. 6 (chest gland marking); COLOMBIA:
Caquetd (Rio Peneya); ecology; behavior (88 day study between August 1975
and February 1976); reidentification of Saguinus fuscicollis, Izawa (1975, 1976).

Saguinus nigricollis cf. graellsiHershkovitz, 1977:1021; COLOMBIA: Caqueta
(Rio Peneya); reidentification of ““Saguinus fuscicollis,’’ Izawa (1975, 1976).

Holotype.—Adult female, flat untanned skin only with hands and feet missing,
Field Museum of Natural History no. 123380; collected 1974 by Tsuyoshe Wa-
tanabe (original number 23), at Rio Peneya, a small tributary of the Rio Caqueta
entering from left (north) about 15 km above mouth of Rio Caguan and about 50
km in straight line below village of La Tagua, Intendencia de Caqueta, Colombia;
altitude approximately 150 m above sea level.

Paratypes.—Two flat untanned skins with hands and feet missing, Field Mu-
seum of Natural History nos. 123379 36, 123381 2, collected 1974 at Rio Peneya
by T. Watanabe; one round skin, Instituto de Ciencias Naturales Universidad
Nacional de Colombia, Bogota, no. 3572, collected January 1960 at Curiplaya,
Rio Caqueta, by F. Medem and H. Granados.

Distribution.—Eastern Colombia between Rios Caqueta, Caguan and Ortegu-
aza and base of the Cordillera Oriental, Intendencia de Caqueta; altitudinal range,
150 to 500 m above sea level.

Coloration of holotype.—Forehead and crown blackish; mantle, throat and
neck blackish, the hairs with one or two narrow orange subterminal bands, base
of those on neck and throat whitish; temporal and genal regions contrastingly
paler than crown, hairs with terminal half blackish, basal half buffy to nearly
whitish and showing through as pale brown or drab; short hairs surrounding
mouth and nostrils gray; blackish mantle tapering to midback and continuing as
a middorsal band to tail base and along proximal 8-9 cm of tail; remainder of tail
entirely blackish; sides of lower half of back modified agouti, the broad orange
subterminal band of hairs giving a striated or marbled appearance; subterminal
band of lateral fringe hairs paler and narrower; arms, legs, rump, sides of body
_and ventral surface of proximal 8-9 cm of tail finely banded orange agouti, the
individual hairs with two orange bands; dark femoral patch indistinct; wrists and
ankles blackish; hands and feet missing but undoubtedly blackish as in the species;
terminal half of hairs of chest mixed orange agouti, saturate orange or saturate
brownish, of belly mixed orange and blackish; skin of face moderately pigmented;
remaining skin unpigmented.

The four skins of the type series are practically indistinguishable.

Measurements.—Those of holotype, a skin only, taken by collector: Head and
body, 220, tail, 340, hind foot, 63; ear 27; weight, 500 g. Paratypes without mor-
phometric data.

Comparisons .—Saguinus nigricollis hernandezi differs from both S. n. graellsi

650 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

and §. n. nigricollis by a blackish, more or less uninterrupted middorsal band on
lower back and proximal portion of tail, and by mainly orange tone of saturate
hairs or pheomelanin bands of individual agouti hairs. Indication or tendency
toward middorsal band or stripe is not evident in graellsi or nigricollis and the
pheomelanin tone is mainly buffy in graellsi, reddish or mahogany in nigricollis.

Saguinus n. hernandezi more nearly resembles nigricollis than graellsi in its
darker, tapered mantle, more broadly banded or striated sides of lower half of
back, and more saturate, less agouti, eumelanin of arms and underparts. On the
other hand, hernandezi more nearly resembles graellsi than nigricollis in its short-
er mantle, finely banded agouti hind limbs, and more extensive agouti of proximal
portion of tail.

Remarks.—Saguinus nigricollis hernandezi is intermediate between graellsi
and nigricollis with respect to color of all chromogenetic fields except for sepa-
ration of the modified agouti of lower back by a more or less defined eumelanin
middorsal band that continues across proximal portion of tail. In effect, hernan-
dezi can be derived from a primitive buffy agouti graellsi type but has diverged
beyond the point where it could give rise to a nigricollis type. In the latter, the
saturate eumelanin of lower back does not extend posteriorly as a stripe to or
beyond tail base but instead tends to spread laterally across sides of trunk.

The three flat untanned skins of hernandezi from Rio Peneya represent the
tamarins recorded by Izawa (1975, 1976) as Saguinus fuscicollis. The samples
sent to the Field Museum for identification in July 1977 by Dr. Tsuyoshe Wata-
nabe, who collaborated in the field with Dr. Kosei Izawa, were identified by me
(1977:1021) as Saguinus nigricollis near graellsi. Izawa (1978:241) accepted this
determination in his third and final report on the ecology and behavior of the
black-mantle tamarin. The saddle-back tamarin, Saguinus fuscicollis was not seen
by Izawa on the north bank of the Rio Caqueta in the Rio Peneya basin. Differ-
ences in color pattern between S. fuscicollis and the larger S. nigricollis are
shown in Figs. | and 2.

Specimens examined.—Four, the holotype and paratypes.

Subspeciation: A Hypothetical Reconstruction

Phenetic and distributional patterns (Figs. 1, 2, 3) of the subspecies of Saguinus
nigricollis suggest that during an early Recent or Late Pleistocene dry period, the
forest habitat of the ancestral form of the species had contracted from a possible
maximum between the Rios Amazonas-Solimoés and Caqueta-Japura to a strip
along the base of the Cordillera Oriental in northern Ecuador and southern Co-
lombia (cf. van der Hammen 1974). Fragmentation of this forest tract with iso-
lation of the Ecuadorian populations from those in Colombia, could result in the
differentiation of two groups. The Ecuadorian group evidently retained with slight
modification the primitive agouti pattern of the ancestral form, whereas the Co-
lombian isolates became more saturate with dominance of eumelanin on fore-
quarters and pheomelanin on hindquarters, the tail excepted. Secondary sepa-
ration of the Colombian stock into two subgroups gave place to divergence of S.
n. nigricollis and S. n. hernandezi. The latter retains more of the ancestral agouti
pattern particularly on hindquarters, but the blackish middorsal band of lower
back and proximal upper surface of tail is sui generis.

With advent of a warmer and more humid climate, dispersal of the three dif-

VOLUME 95, NUMBER 4 651

Saguinus nigricollis Group

(Ih

S. fuscicollis S. nigricollis

Fig. 2. Facial color patterns of species of Saguinus nigricollis group. Contrasting pale brown
temporal and genal regions, constant features of S. nigricollis, are not present in S. fuscicollis.

ferentiated forms of Saguinus nigricollis does not appear to have coincided pari
passu with spread of the sylvan habitat from its near-cordilleran base to the banks
_ of the Amazonas. Possible explanations are discussed beyond.

Geographic Distribution of the Subspecies of
Saguinus nigricollis

The range of Saguinus nigricollis graellsi in eastern Ecuador and northeastern
Peru extends from the Rio Sucumbios-Putumayo at the northwestern boundary
between Ecuador and Colombia, east to the Rios Guepi and Lagartococha, south
through the Rio Aguarico basin into the right bank basin of the Rio Napo, thence
south from between the river and foothills of the Cordillera Oriental to the north
bank of the Maranon-Amazonas in Peru; altitudinal range is from about 100 m to
600 m or more above sea level.

All locality records of museum-preserved specimens lie north of the disputed
boundary between Ecuador and Peru (Fig. 3) except for a point opposite Tara-
poto, Rio Napo (4), the mouth of the Rio Curaray (4), and Destacamento (now
Francisco Orellana) at the mouth of the Rio Napo (5).

As hypothesized in the preceding section, Saguinus nigricollis graellsi dis-
persed to the east and south as its forest refuge expanded from the cordilleran
base in northern Ecuador to the Maranon-Amazonas. Primary dispersal routes
were forests spreading along the banks of the Rios Napo, Tigre and Pastaza.
Sanguinus n. graellsi gained the Rio Amazonas via the right bank of the Rio

_Napo but is otherwise unknown in Peru. However, scientific studies or collections

of mammals have not been reported from the northern halves of the Rios Tigre »
and Pastaza basins in the territory between the Maranon and the Ecuadorian-
Peruvian boundary.

The geographic range of graellsi shown in Fig. 3 as questionably extending to
the north bank of the Maranon is presumptive. The superficially similar Callimico
goeldii was unknown on the north bank until 1970. The earliest published record

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

652

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VOLUME 95, NUMBER 4 653

for Colombia is 1968, and Callimico has yet to be recorded from Ecuador where
it surely occurs. For history and distribution of Callimico, see Hershkovitz (1977:
864, 892, 904).

If cryptic habits do not explain the apparent absence of Saguinus nigricollis
graellsi west of the Rio Napo basin in Peru, perhaps lack of suitable habitat not
previously occupied by potential competitors is the answer. Much of the region
between the lower Rios Napo and Pastaza is black water swamp and floodland
where the slightly smaller saddle-back tamarin, Saguinus fuscicollis lagonotus,
abounds. This representative of the most wide-spread callitrichid species can be
derived from an ancestral stock resembling Saguinus fuscicollis illigeri of the
south bank of the Maranon by a modification of the mantle from primitive agouti
to saturate pheomelanin. Presumably, the /agonotus founder colony became iso-
lated from the i/ligeri-like parental form by a river bend cutoff with passive trans-
fer to the north bank of the Maranon. Northern dispersal of the differentiated S.
fuscicollis lagonotus may have excluded S. nigricollis graellsi from the lower
halves of river basins shared in their upper halves by both species but where
habitats are particularly favorable for S. n. graellsi.

Preserved specimens of Saguinus nigricollis graellsi from Colombia are un-
known to me. Reports of graellsi seen north of the Ecuadorian border (see Hersh-
kovitz 1977:629, 1021 for discussion) may refer to another kind of tamarin, per-
haps Saguinus nigricollis nigricollis or S. n. hernandezi.

Saguinus nigricollis nigricollis has been known to inhabit the region between
the Rio Amazonas-Solimoés and the Rio Putumayo-I¢a from the mouth of the
latter in Brazil to the mouth of the Rio Napo in Peru. The geographic range,
however, probably comprises the entire region between the Rios Napo and Pu-
tumayo in Peru upstream to the Rio Lagartococha-Aguarico at the Ecuadorian-
Peruvian border. It may also include the interfluvial basin between the Rios Pu-
tumayo-I¢a and Caqueta-Japura from the north shore of the Amazonas-Solimoés
to near the base of the Cordillera Oriental in Colombia. The altitudinal range is
about 90 m to 400 m above sea level.

Delineation of the presumptive maximum range of Saguinus nigricollis nigri-
collis derives solely from the skin of a young individual labelled ‘‘“Caqueta-Pu-
tumayo,’’ an area identified in Fig. 3 by vertical hatching with an interrogation
sign on a white circle with arrows pointing in three directions. The specimen,
collected “January 1960,’ by H. Granados and H. Arévalo, is indistinguishable
from Rio Amazonas-Solimoés Saguinus nigricollis nigricollis. These data prove
that the geographic range of S. n. nigricollis in the upper half of the watershed
embraced by the Rios Putumayo and Caqueta closes the geographic gap between
the range of S. n. graellsi and S. n. hernandezi. The locality record “‘Caqueta-
Putumayo”’ also indicates that, as far as known, upstream populations of nigri-
collis are isolated from downstream sister populations between the lower Rio
Putumayo-Ic¢ca and Rio Amazonas-Solimoés. Had Granados and Arévalo taken
Spix’s black-mantled tamarin on the right bank of the Rio Putumayo where on
‘January 1960” they also collected Callimico goeldii at Quebrada del Hacha
(Hershkovitz 1977:926), the range of S. n. nigricollis would not appear disjunct.
This hypothetical eventuality would leave the range of S. n. hernandezi discon-
tinuous with respect to those of graellsi and nigricollis.

654 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

The authenticity of the ‘‘Caqueta-Putumayo”’ record is not questioned but it
may be premature to extrapolate the enormous extension of the geographic range
of Saguinus nigricollis nigricollis indicated by this datum alone. This species has
not been collected or observed between the Rios Putumayo and Caqueta by others
including Humboldt, Lako and Salachi, Erikson, Hershkovitz, Moynihan, and
Izawa (Hershkovitz 1977:643, 926, 932, 1038). Watanabe collected only the sad-
dle-back tamarin, Saguinus fuscicollis fuscus between the Rios Putumayo-I¢a
and Caqueta-Japura (actually, right bank Rio Caqueta and El Cedral above Puerto
Asis, specimens in the Field Museum).

The range of Saguinus nigricollis nigricollis (Fig. 3) is separated from that of
S.n. graellsi by the Rio Napo and its tributary the Rio Lagartococha-Aguarico,
and from that of S. n. hernandezi by the upper Rio Caqueta. Saguinus n. ni-
gricollis is parapatric with the saddle-back tamarin S. fuscicollis nigrifrons along
the Rio Amazonas and S. fuscicollis fuscicollis along the Solimoés. It is sympatric
with Saguinus fuscicollis tripartitus between the Rios Napo and Putumayo (Puer-
to Narino, Rio Amazonas, Hernandez Camacho and Cooper 1976:39). If the range
of nigricollis extends to the Rio Caqueta, then it is also sympatric with S. fus-
cicollis fuscus and parapatric with the mottled-face tamarin Saguinus inustus
along the Caqueta between the Rios Caguan and Apaporis (see below).

Saguinus nigricollis hernandezi occurs between the Rios Caqueta, Caguan,
Orteguaza and base of the Cordillera Oriental in the Intendencia de Caqueta,
eastern Colombia. The range very likely includes the segment between the Rios
Caqueta and Orteguaza. Altitudinal range is from about 150 m to 500 m above
sea level.

Saguinus nigricollis hernandezi is sympatric with the saddle-back tamarin, S.
fuscicollis fuscus, parapatric with S. n. nigricollis along the Rio Caqueta and
possibly with the mottled-face tamarin S. inustus along the Rio Caguan. Accord-
ing to Thomas R. Defler (personal communication, March 1981), S. inustus occurs
in the region between the Rios Apaporis and Caqueta, but his records are of
individuals seen and collected along the Rio Miritiparana, a left bank tributary of
the Rio Caqueta near the Brazilian border. The range of Saguinus inustus possibly
extends west to the Rio Caguan.

Attribution and Acknowledgments

The northernmost race of the black-mantle tamarin is named in honor of Dr.
Jorge Hernandez Camacho, one of Colombia’s most respected biologists and
perhaps one of the last of South America’s authentic naturalists. Investigators of
Colombian biota have found Jorge’s expertise in their particular discipline, no
matter which, an enormous asset unstintingly placed at their service and much
of their success is owed to his advice, guidance and practical assistance in both
his private and official capacities. |

I take this occasion to express my gratitude to Dr. Hernandez for a critical
reading of page proof of most of the text of volume 1 of my (1977) Living New
World Monkeys. The value of his contributions to the accuracy, clarity and ob-
jectives of the book are inestimable. It seems, however, that my note of appre-
ciation on page xiii of page proof of the already locked in “‘Introduction’’ to the
volume (see Index, p. 1056) was deleted for replacement by the later submitted
‘*Corrigenda.”’

VOLUME 95, NUMBER 4 655

The three Rio Peneya skins described here as Saguinus nigricollis hernandezi
were received by the Field Museum of Natural History from Tsuyoshe Watanabe
of the Primate Research Institute, Inuyama, Japan, in exchange for the identifi-
cations. Loan of the paratype of S. nigricollis hernandezi (ICN 3572) and the skin
of S. nigricollis nigricollis (CN 3556) from *‘Caqueta-Putumayo’’ by authorities
of the Instituto de Ciencias Naturales of Bogota, is acknowledged with thanks.
Figure | was drawn by Field Museum staff artist Marlene Warner. Technical
assistance and typing of the manuscript was provided by Barbara Brown.

Preparation of this report was aided in part by a grant (RRO1226) from the
National Institutes of Health.

Literature Cited

Hernandez Camacho, Jorge, and Robert W. Cooper. 1976. The nonhuman primates of Colombia.—
In R. W. Thorington, Jr. and P. G. Heltne (eds.), Neotropical primates. Field studies and
conservation.—National Academy of Sciences, Washington DC, pp. 35-69, 13 figs.

Hershkovitz, Philip. 1966. Taxonomic notes on tamarins, genus Saguinus (Callithricidae, Primates),

with descriptions of four new forms.—Folia Primatologia 4:381—395, 4 figures.

1977. Living New World Monkeys (Platyrrhini) with an introduction to primates. Volume

1. University of Chicago Press, Chicago, pp. xiv + 1117, 520 figs., 7 color plates.

Izawa, Kosei. 1975. Foods and feeding behavior of monkeys in the upper Amazon basin.—Primates
16(3):295-3 16, 8 figures.

—. 1976. Group sizes and composition of monkeys in the upper Amazon basin.—Primates

17(3):367-399, 6 figures.

1978. A field study of the ecology and behavior of the black-mantle tamarin (Saguinus
nigricollis).—Primates 19(2):241—274, 9 figures.

van der Hammen, T. 1974. The Pleistocene changes of vegetation and climate in tropical South
America.—Journal of Biogeography 1:3—26, 21 figures.

Field Museum of Natural History, Roosevelt Road at Lake Shore Drive, Chi-
cago, Illinois 60605.

Gazetteer of Collecting Localities of Saguinus nigricollis
Shown in Fig. 3

COLOMBIA
Caqueta—Saguinus nigricollis hernandezi
1. Curiplaya, Rio Caqueta, 0°16’N, 74°52’W. F. Medem and H. Granados, January 1960.
2. Peneya (Rio), mouth N side Rio Caqueta, 0°07'S, 74°22’W—type locality. Observations by
K. Izawa at Puerto Japon, 0°7'’N, 74°24'W, and Puerto Tokio, 0°12'N, 74°23’W, August 1975—
February 1976. Collection by T. Watanabe, 1974.
Amazonas—Saguinus nigricollis nigricollis
3. Leticia, Rio Amazonas, 4°15’S, 69°56’W. J. N. Layne, February 1956 at 100 m. G. Erikson
(purchased).
‘“Caqueta-Putumayo’’—Saguinus nigricollis nigricollis
? Locality between Rios Caqueta and Putumayo where boundaries of the two Intendencias
of the same names approach or could not be determined at point of collection. H. Granados
and H. Arévalo, January 1960.

PERU
Loreto—S.n. graellsi and S. n. nigricollis
4. Tarapoto (opposite), Rio Napo, 2°15’S, 74°06’'W—type locality of Saguinus nigricollis graell-
si. M. Jiménez de la Espada, June, July 1865.
4. Curaray (Rio), boca (=mouth) at Rio Napo, 2°22'S, 74°05’'W—Saguinus nigricollis graellsi.
Olalla and Sons, October, November, December 1925 at 140 m.

656

Ke)

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

. Destacamento (=Francisco Orellana), Rio Napo, at junction with Rio Amazonas, 3°21’ =

72°45'W—Saguinus nigricollis graellsi. M. Jiménez de la Espada, 1865.

. Apayacu (Rio), mouth at Rio Amazonas, 3°21'S, 72°07'W, at 100 m—Saguinus nigricollis

nigricollis. Olalla and Sons, December 1926, January 1927.

. Ampiyacu, Rio Amazonas, 3°19’S, 71°51’W—Saguinus nigricollis nigricollis. C. Freese, Oc-

tober, November, 1974 (sight records).

. Pebas, Rio Amazonas, 3°10’S, 71°48’W—Saguinus nigricollis nigricollis. Castelnau and De-

ville, about 1847; Olalla and Sons, October, 1926; R. W. Hendee, January 1928 at 91 m.

. Chimbote (opposite), Rio Amazonas, 3°49’S, 70°41’W—Saguinus nigricollis nigricollis.
. Yahuas Territory, see Santa Maria de Las Yaguas.
. Santa Maria de Las Yaguas, near Rio Putumayo at Colombian border, 2°43'S, 69°58’W—

Saguinus nigricollis nigricollis. J. J. Mounsey, August 1913.

BRAZIL—Saguinus nigricollis nigricollis
Amazonas

10.
I,

Ze

Santa Rita, Rio Solimoés, 3°30’S, 69°20’W. W. Ehrhardt, September 1926.

Sao Paulo de Olivenca, Rio Solimoés, 3°27’S, 68°48’W (type locality). Spix and Martius
Expedition.

Between Rio Solimoés and Rio I¢a. Spix and Martius Expedition.

ECUADOR—Saguinus nigricollis graellsi

Napo
3.

14.
IDs

16.
LW:
We
18.

Santa Rosa de Sucumbios, Rio San Miguel, right bank tributary of Rio Putuamayo, 0°15’N,
76°27'W. Olalla, November 1931.

San Francisco, Rio Napo, 0°30’S, 76°22'W. P. Hershkovitz, February, March, 1936, at 200 m.
Lagarto Cocha (Rio), mouth at Rio Aguarico, 0°39’S, 75°16’W. Olalla and Sons, January
1926, at 190 m.

Sumaco, 0°34’S, 77°30'W. Olalla and Sons, December 1923, January 1924.

Avila, 0°38’S, 77°25’W, at 600 m. W. Clark-McIntyre, May 1937; A. Olaila, September 1938.
San José (below), Rio Suno, 0°31'S, 77°25'W. Olalla and Sons, March, April 1924. |
Suno (Rio), 0°42'S, 77°08’W. Olalla Brothers, 1923, March 1924, February 1929.

Pastaza

19.

20.

Alle

Yana Rumi (Rio), mouth at Rio Pindo Yacu, 1°38’S, 76°59’W. R. Olalla, December 1934,
February 1935.

Pindo Yacu (Rio), joins Rio Cunambo, upper Rio Tigre at 2°08’S, 76°04’ W. R. Olalla, October,
1934, above junction with Rio Cunambo at 250 m.

Pastaza (Rio), 2°05'S, 500 m. C. S. Webb.

22. Capahuara or Capihuara (Rio), mouth at Rio Pastaza, 2°3’S, 76°51’W. R. Olalla, November

1934, above mouth at 300 m.

PROC. BIOL. SOC. WASH.
95(4), 1982, pp. 657-687

REVIEW OF THE GENUS MICROGNATHUS DUNCKER
(PISCES: SYNGNATHIDAE), WITH DESCRIPTION
OF M. NATANS, N. SP.

C. E. Dawson

Abstract.—The urophorine (tail-pouch) pipefish genus Micrognathus Duncker
is reviewed and four Indo-Pacific and one Atlantic species are recognized. Other
nominal species usually included in Micrognathus are provisionally referred to
the genus Halicampus Kaup. Micrognathus natans n. sp., characterized in part
by a modal count of 14 trunk rings, is described from the Philippines, eastern
Australia, New Caledonia, and the Fiji Islands. Other Indo-Pacific species include
the type-species, M. brevirostris Ruppell, sensu Kaup, 1856), M. andersonii
(Bleeker), and M. micronotopterus (Fowler). Two of these (andersonii, micro-
notopterus) typically lack protruding, hook-like, posterior angles on the tail rings,
whereas rings on the distal half or more of the tail have prominent hook-like
spines or points in M. brevirostris. Micrognathus andersonii (modally 16 trunk
rings) ranges from the northern Red Sea to Japan (Honshu Is.), Samoa and Tonga,
whereas M. micronotopterus (modally 15 trunk rings) is known from northern
and western Australia, western Indonesia, Singapore and the Philippines. Mi-
_crognathus brevirostris includes two subspecies, M. b. brevirostris in the Red
Sea and M. b. pygmaeus in NE Australia, eastern Indonesia, Marshall Is. (Bikini)
and Tahiti. These subspecies share a modal count of 15 trunk rings, but differ in
preserved coloration, as well as in numbers of pectoral-fin rays and subdorsal tail
rings (respectively, 9-11 and 3.75—4.75 versus 10-13 and 2.75-4.0 in pygmaeus).
There are no confirmed Indian Ocean records of any species of Micrognathus
between Madagascar, the SE tip of India and the NW coast of Australia. The
single Atlantic species (M. erugatus), characterized by 20 trunk rings, is known
only from the Brazilian type locality. A key is provided, each taxon is diagnosed
and illustrated, and maps delineate distributions of Indo-Pacific species as deter-
mined from materials examined.

The genus Micrognathus Duncker, 1912 has never been differentiated ade-
quately from other urophorine (tail-pouch) pipefish genera with the same config-
uration of principal body ridges (e.g. Halicampus Kaup, Trachyrhamphus Kaup),
and has never been subject to critical taxonomic study. Herald and Randall (1972)
included three subgenera and some 15 species in Micrognathus, and three addi-
tional species have since been described from Atlantic and Pacific waters (Herald
and Dawson 1974, Dawson and Allen 1981, Fritzsche 1981). Many nominal species
lack an adequate diagnosis or description, available figures are often poor, and
correct identification of individual specimens is often difficult or impossible. In
fact, the identity of the type-species (Syngnathus brevirostris Ruppell) is uncer-
tain, and at least three taxa are commonly identified in collections and the
literature as Micrognathus brevirostris. This review, based on examination of
available museum material, recognizes four species of Micrognathus in the Indo-

658 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Pacific region and one species (M. erugatus Herald and Dawson) in the western
Atlantic Ocean. One Indo-Pacific species is described as new, two [M. andersonii
(Bleeker), M. micronotopterus (Fowler)] are resurrected from synonymy, and M.
brevirostris 1s considered to include one subspecies in the Red Sea and another
in the western Pacific Ocean.

Methods and Materials

Measurements are in millimeters (mm) and some are referred to standard length
(SL), total length (TL) or head length (HL). Counts of trunk rings begin with that
bearing the pectoral fins and end with that bearing the anus (anal ring); all fin-
rays are counted separately; subdorsal rings are estimated to the nearest 1/4-ring
interval before (+) and after (—) the anterior margin of the Ist tail ring (O0-point).
As employed here, the term “‘venter’’ refers to the ventral surface of head or
body; color descriptions are from specimens preserved in alcohol; other methods
are those of Dawson (1977). Care must be taken in counting the number of trunk
rings, since any error may result in misidentification. Materials examined are
listed by general localities from west to east and roughly north to south; depths
are in meters (m); maps delineate general localities and each symbol represents
one or more collections.

Abbreviations for repositories of materials examined are: AMNH—American
Museum of Natural History, New York; AMS—Australian Museum, Sydney;
ANSP—Academy of Natural Sciences of Philadelphia; BMNH—British Museum
(Natural History), London; BPBM—Bernice P. Bishop Museum, Honolulu; CAS—
California Academy of Sciences, San Francisco; CAS-SU—Stanford University
collections, now at CAS; CSIRO—CSIRO New South Wales Fisheries Labora-
tory, Cronulla; FMNH—Field Museum of Natural History, Chicago; GCRL—
Gulf Coast Research Laboratory Museum; HUJ—Hebrew University of Jerusa-
lem; KFRS—Kanudi Fisheries Research Station, Papua New Guinea; MCZ—
Museum of Comparative Zoology, Harvard University, Cambridge; MNHN—
Museum National d’Histoire Naturelle, Paris; MZUSP—Museu Paulista, Uni-
versidade do Sao Paulo; NMW—Naturhistorisches Museum, Vienna; NTIM—
Northern Territory Museum, Darwin; QM—Queensland Museum, Brisbane;
RMNH—Riksmuseum van Natuurlijke Historie, Leiden; RUSI—J. L. B. Smith
Institute of Ichthyology, Grahamstown; SAM—South African Museum, Cape
Town; SIlO—Scripps Institution of Oceanography, La Jolla; SMF—Natur-Mu-
seum und Forschungs-Institut Senckenberg, Frankfurt am Main; TAU—Tel-Aviv
University; UG—University of Guam, Agana; UMMZ—Museum of Zoology,
University of Michigan, Ann. Arbor; USNM—National Museum of Natural His-
tory, Smithsonian Institution, Washington, D.C.; UWCF—University of Wash-
ington College of Fisheries, Seattle; WAM—Western Australian Museum, Perth;
Y CM—Yokosuka City Museum; ZMA—Zoologisch Museum, Amsterdam; ZMB—
Zoologisches Museum, Museum fir Naturkunde der Humboldt-Universitat, Ber-
lin; ZMUC—Zoologisk Museum, University of Copenhagen.

Micrognathus Duncker

Micrognathus Duncker, 1912:235 (type-species, Syngnathus brevirostris Ruppell
1838, by original designation).

VOLUME 95, NUMBER 4 659

Diagnosis.—Superior trunk and tail ridges discontinuous near rear of dorsal-
fin base, not arched strongly dorsad below dorsal-fin base in subadults-adults;
inferior trunk ridge and lateral tail ridge end near anal ring; lateral trunk ridge
confluent with inferior tail ridge; venter of trunk V-shaped, the median ridge
distinct but not enlarged or Keel-like. Median dorsal snout ridge low, entire,
somewhat concave in lateral profile, originating on anterior half of snout, ending
on or before middle of interorbital, not confluent with orbital ridges; lateral snout
ridge absent; dorsal rim of orbit somewhat elevated; interorbital concave or a
little depressed; prenuchal, nuchal and frontal ridges low, arcuate, essentially
entire; principal (longitudinal) opercular ridge complete in young, usually incom-
plete or infrequently obsolete in adults, ridge straight or angled upward toward
gill opening; other head ridges low, mainly entire; head without spines, knobs,
denticules or serrations. Principal trunk ridges low to somewhat elevated, without
prominent indentations or notches between anterior and posterior margins of
rings; dorsum often a little depressed between superior trunk ridges; tail rect-
angular in section or with principal ridges angled progressively laterad toward
caudal fin and with surfaces depressed or concave between; ridges notched or
indented between rings; posterior angles of tail rings not produced strongly (Figs.
la, b) or forming prominent hook-like projections (Fig. lc); ridge margins entire,
granular or rough, never clearly denticulate or serrate; scutella without longitu-
dinal keels; dermal flaps absent from eye, usually present on head and/or body,
generally small, never greatly enlarged or profusely branched; dorsal-fin origin
between anterior margin of penultimate trunk ring and posterior margin of Ist tail
ring, fin-base not clearly elevated in subadults-adults; pectoral and caudal fins
small, rounded; anal fin present. Trunk rings 14—20, total rings 41-56, total sub-
dorsal rings 3.25—5.75, dorsal-fin rays 16-24, pectoral-fin rays 9-15, anal-fin rays
2-4, caudal-fin rays typically 10. Head length 6.9-11.3 in SL, snout length 2.2-
4.0 in HL, length of dorsal-fin base 0.9-1.9 in HL (data from subadults-adults).
Brood pouch located below anterior 10—18 tail rings; pouch plates little enlarged;
pouch folds present; pouch closure the everted type of Herald (1959); brood-pouch
eggs deposited in 1-2 layers and in 1-7 transverse rows. Without odontoid pro-
cesses in jaws (Dawson and Fritzsche 1975) or bony inclusions in gill membranes
(Dawson 1978); all branchial elements, except 3rd basibranchial, present (Fritzsche
1980).

Comparisons.—Among urophorine (tail-pouch) pipefish genera, the principal
body ridge configuration of Micrognathus and combined presence of a 10-rayed
caudal fin, an anal fin, a brood pouch with both pouch plates and folds, and a
non-prehensile tail are shared only with Halicampus Kaup. These taxa differ
principally in the development and ornamentation of head and body ridges, and
in the type and development of dermal flaps. Micrognathus is in part character-
ized by a low, entire, median dorsal snout ridge, absence of a lateral snout ridge,
absence of strongly elevated head or trunk ridges, and absence of spines, denti-
cules or serrations on the head or trunk. In contrast, species of Halicampus have
an elevated and/or spiny median snout ridge, the lateral snout ridge or spine is
usually present, spines are often present elsewhere on the head, and the head
and trunk ridges are often elevated strongly with denticulate or serrate margins.
In Micrognathus, dermal flaps are absent from the eye and are elsewhere usually
small, slender, and little branched. Dermal flaps are present on the eye of most

660 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Cross-sectional and oblique-longitudinal aspects of distal portion of the tail in species of
Micrognathus. a—b: M. andersonii, M. erugatus, M. micronotopterus, M. natans; c: M. brevirostris.

species of Halicampus and those on the head and body are often large, sometimes
spatulate, and frequently multi-branched. Duncker (1915) noted that the dorsal-
fin base is low in Micrognathus and elevated in Halicampus, but this appears to
be a shared character, albeit less obvious in Micrognathus. In Micrognathus,
young fish may have a somewhat elevated dorsal-fin base, and young and some
adults may have the superior trunk ridge arched a little dorsad below the dorsal-
fin base.

Remarks.—Herald (1953) included two subgenera, Anarchopterus Hubbs, 1935
(type-species, Siphostoma crinigerum Bean and Dresel) without anal fin and Mi-
crognathus (with anal fin), in the genus Micrognathus. Subsequently, Herald and
Randall (1972) introduced Minyichthys (type-species, Micrognathus brachyrhinus
Herald) as a third subgenus, differentiated only by high numbers of trunk rings
(19-21 versus 13-17), a ‘“‘very short snout’’ and small size at maturity (<50 mm
SL). Dawson and Allen (1981) restored generic rank to Anarchopterus, and Daw-
son (1982) treated Minyichthys as a genus, characterized in part by an 8-rayed
caudal fin. Species included in the remaining subgenus (Micrognathus, sensu
Herald and Randall 1972) share a high degree of consistency in numbers of trunk
rings (maximum range of 3 for any species), but exhibit considerable variation in
gross morphology and include at least two lineages of somewhat similar pipefish-
es. One of these, represented by the genus Micrognathus, includes the species
treated here. All other species, including the western Atlantic M. crinitus (Jenyns,
1842), M. spinirostris Dawson and Allen, 1981, and the remaining Indo-Pacific
members of Herald’s (1953) subgenus Micrognathus, are provisionally referred
to the genus Halicampus Kaup. Micrognathus and Halicampus are similar in
many respects and may prove to differ only at the subgeneric level. However, I

VOLUME 95, NUMBER 4 661

retain generic rank for Micrognathus pending completion of studies on Halicam-
pus.

Although problems exist with the identity of the type-species of Micrognathus
(see Remarks under M. b. brevirostris), common usage argues strongly for pres-
ervation of Duncker’s genus name, and Syngnathus brevirostris Riippell (sensu
Kaup 1856) is provisionally accepted as the type-species of Micrognathus. In
addition to sharing features diagnosed above, these pipefishes are small (probably
seldom exceeding 80 mm SL), some are locally abundant, and all but one species
(natans) are known to occur in tidepools and shallow reef or shore habitats.
Distribution of Indo-Pacific species extends from the northern Red Sea to Japan
(Honshu Is.), the Marshall Is. (Bikini) and Tahiti. However, within the Indian
Ocean, the genus is known only from the eastern coast of Africa, Madagascar,
the southeastern tip of India, western Indonesia and northwestern Australia.

Key to the species and subspecies of Micrognathus

Peminunk mnmesl4—17 (Indo-Pacihic) 14.50. ccc sade nis dios alegethe'e sue ee wa was yy
— Trunk rings 20 (western Atlantic) ........... erugatus Herald and Dawson

2. Trunk rings usually (in 93% of specimens examined) 14, snout depth in
ponoutslemoth2:9=4,0:GQ = 3.4). oc aed ewe ce volcanoes natans Nn. sp.

— Trunk rings usually (in 99.9%) 15-17, snout depth in snout length 1.6-3.1
} (Gms Mew AGO) ae gatnde pray syrc, Ayes on ood tas ob serensbedis: osicuetras a Stale eToys. Acasmnsvain. Sets ORY oun Bumesbe 3
nem s: 14— Oil Si1m 99%) cost, Pe a. SEB otras a4 8 Ss yea haren ss 4
lnonke nies, 1S—17 (6-17 in 99.77%). ok ds eden es andersonii (Bleeker)

4. Principal tail ridges angled laterad (Fig. Ic), dorsal-fin rays 18-21 (19-21
imo) snout lensth averages 3.0-3.l in HL... 2... ee le oe ee 5

— Principal tail ridges not angled laterad (Fig. la), dorsal-fin rays 17-19 (17—
18 in 94%), snout length averages 2.6 in HL ..... micronotopterus (Fowler)

5. Subdorsal tail rings 3.75—4.75 (4.0 or more in 85%); pectoral-fin rays 9-

11; opercle without minute dark spots or ocellus, often with narrow pale
IDES (RECISS2) Reyes ere aa een brevirostris brevirostris (Ruppell)

— Subdorsal tail rings 2.75—4.25 (3.75 or fewer in 92%); pectoral-fin rays 10-

13; opercle without narrow pale bars, often with pale-margined ocellus

and/or minute dark spots in subadults and adults (W Pacific)...........
esi te Pid vb ace lache aaa buoya'eyors brevirostris pygmaeus Fritzsche

Micrognathus brevirostris (Ruppell)

Diagnosis.—TYrunk rings modally 15, principal ridges angled laterad on distal
half of tail, dorsal-fin rays 18-21 (usually 19-20), snout length averages 3.0—3.1
in HL.

Description.—See subspecies.

Comparisons.—Characters in key and diagnosis distinguish M. brevirostris from
congeners. Additionally, M. brevirostris lacks dark blotches above the lateral
trunk ridge (often present in M. andersonii and M. micronotopterus) and has a
higher average HL in SL ratio than M. natans (ca. 9.4 versus 7.4).

Remarks.—As interpreted here, M. brevirostris includes one subspecies in the
Red Sea and another in the western Pacific Ocean. These are distinguished by

662 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

differences in some meristic and proportional values, as well as in preserved
coloration (see key and diagnoses of subspecies).

Micrognathus brevirostris brevirostris (Ruppell)
Figs. 2-4

Syngnathus brevirostris Ruppell, 1838:114 (orig. descr.; Massaua, Red Sea).

Diagnosis.—Subdorsal tail rings usually 4.0-4.75, pectoral-fin rays usually 10,
without minute dark spots on opercle, side of head often with irregular narrow
bars. |
Description.—Rings 14-16 + 28-31 (15 + 28-31 in 95%), dorsal-fin rays 18-
21, pectoral-fin rays 9-11 (modally 10), anal-fin rays usually 3, subdorsal rings
1.25-0.25 + 3.75-4.75 = 4.25-5.5 (see Tables 1—3 for additional counts). Propor-
tional data, based on 19 specimens 36.0-62.0 (« = 45.8) mm SL follow: HL in
SL 8.7-11.3 (9.7), snout length in HL 2.9-4.0 (3.1), snout depth in snout length
1.6—2.3 (2.0), length of dorsal-fin base in HL 1.0-1.2 (1.1), pectoral-fin length in
HL 3.7-6.0 (4.5), length of pectoral-fin base in pectoral-fin length 1.2—2.0 (1.7),
anal ring depth in HL 2.0-2.6 (2.3), trunk depth in HL 1.6—2.1 (1.9). Longitudinal
opercular ridge essentially complete in larvae and juveniles, incomplete (Fig. 2)
or occasionally obsolete in adults. Distal half or more of tail with principal ridges
angled or flared laterad and with posterior angles of rings produced to form hook-
like points (Fig. 1c); dermal flaps usually short, typically slender, usually simple
but sometimes with 1—2 subterminal branches or bifurcate distally.

Ground color pale to dark brown in subadults—adults; both pale and dark spec-
imens may occur in a single collection, irrespective of sex or state of maturity;
eye usually with indications of dark bars radiating from pupil. Pale fish usually
have some brownish shading on side and venter of head and on lower half of
trunk; some have head and body mottled with tan; fins mainly hyaline. Dark
specimens may be plain or peppered with minute pale spots, dorsal and pectoral
fins are brownish and caudal fin is brown with pale distal margin. Among 23
recently preserved fish, 19 (83%) have diffuse dark spots on or near the lateral,
inferior and median ventral ridges of the anterior 6—12 trunk rings. Arrangement
of spots, a vertical row of patches of dark microchromatophores, suggests that
the lower half of the trunk is barred in life. Additionally, recently preserved
specimens often have several to many narrow, irregular, pale bars or lines on the
side of the snout, suborbital and opercle; these markings may extend above the
opercle and on the venter of the head.

Comparisons.—Micrognathus b. brevirostris differs from M. b. pygmaeus in
having fewer pectoral-fin rays (modally, 10 versus 11—12) and in having a higher
average number of subdorsal tail rings (4.2 versus 3.4). These subspecies evi-
dently share the presence of bars on the lower half of the trunk, but M. b.
brevirostris lacks dark spots on the opercle (usually present in M. b. pygmaeus
>30 mm SL) and narrow pale bars on the head (present in many M. b. breviros-
tris) have not been found in examined specimens of M. b. pygmaeus. These
pipefishes are closely related, but the differences noted here and in descriptions,
together with allopatric distributions, are sufficient for recognition as subspecies.

Remarks.—Although the above-described taxon was identified with Syngna-

VOLUME 95, NUMBER 4 663

ou

Fig. 2. Micrognathus brevirostris brevirostris. Lateral and dorsal aspects of head and anterior
trunk rings, together with section of body illustrating ridge configuration and dorsal and anal fins.
From 51.5 mm SL, male (BMNH 1860.11.9.60).

thus brevirostris Ruppell by Kaup (1856), and subsequent workers have accepted
Kaup’s interpretation, the identity of Ruppell’s species is uncertain. Ruppell’s
(1838) unillustrated original description may be roughly translated:

Head measures a tenth of the body length and the snout equals a third of
the head length; opercle and vertex smooth; the body, although 7-angled,
appears cylindrical; there are 18 ring segments between head and anus, 11
along the brood pouch, and 19 in the remainder of the tail; the short caudal
fin is rounded; pectoral fin 7; dorsal fin 19; caudal fin 7; body grayish yellow

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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VOLUME 95, NUMBER 4 665

~ Table 3.—Frequency distributions of trunk, tail, and total subdorsal rings in species of Micro-
gnathus. Plus (+) indicates dorsal-fin origin in advance of 0-point (anterior margin of Ist tail ring);
minus (—) indicates origin on tail. * Designates holotype.

Subdorsal trunk rings

Species ap WS) 1.50 1.25 1.00 0.75 0.50 0.25 0.00 0.25 0.50 —

b. brevirostris py) 7 11 15 6

b. pygmaeus 2 2 34* ZT 8 3

andersonii 9 22 =D) 151 76* 33 3

micronotopterus l 3) 16* 12 4 |

natans 3) 13 31 11* 5
erugatus 1s

Subdorsal tail rings

LOA DS QI SATO BS SSN ISS ac 4.25 Yb \1)
b. brevirostris 6 13 15 4 3
b. pygmaeus 3 8 DS 24 10 Se 1
andersonii B 10 31 47 63 58 61 42* 27 5 y)
micronotopterus 2 6* D 11 8 > 3
natans 2 4 14* 15 20 i 1
erugaius li

Total subdorsal rings

3 25pe B50 tr Berd 4.00 AOS ASO GIS 5.00 5125 5.50 5.75
b. brevirostris 5 6 12 al 6 1
b. pygmaeus 3 23 28 13 8 1s
andersonii 5) jap 63 59 78 70* 38 11 3
micronotopterus D 4* 9 13 6 3
natans ] i 17 DD, 9 1
erugatus 1a

in alcohol, with numerous lighter spots; edges of body rings unspotted and
appear darker; head with some whitish irregular bands; fins dark gray; body
length 2.5 inches, from Massaua.

Ruppell’s remarks concerning body proportions, opercle, vertex, 7-angled body,
brood pouch, size at maturity, size and shape of caudal fin, and coloration are
not diagnostic and could apply to many pipefishes. According to modern methods,
occurrence of 18 rings between the head and the anus would result in a count of
19 or 20 trunk rings, but no Red Sea species historically referred to Micrognathus
has more than 17 trunk rings. Similarly, pectoral-fin rays number 9-15 in all Indo-
Pacific taxa previously referred to Micrognathus, 19 dorsal-fin rays occur in sev-
eral Red Sea pipefishes, and 7 caudal-fin rays is an atypical number for any
pipefish. No clue is provided to the configuration of the principal body ridges,
and, aside from reference to the male, there is no indication of the number of
specimens obtained by Ruppell. Subsequently, Kaup (1853, 1856) referred an
unstated number of specimens, collected by Ruppell but differing significantly
from the foregoing description, to this species (as Corythoichthys brevirostris). |
have found no evidence to indicate that the specimens treated by Kaup are con-
specific with Ruppell’s specimen(s) of Syngnathus brevirostris.

666 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Micrognathus brevirostris brevirostris. Top, BMNH 1860.11.9.60 (51.5 mm SL male).
Middle and bottom, GCRL 18154 (43.5 mm SL male, and 48 mm SL female).

Since Ruppell’s (1838) description fails to define any character or character
combination diagnostic of any known pipefish, the name brevirostris could well
be treated as a nomen dubium. This, however, would introduce a number of
undesirable nomenclatural problems. Among these would be the suppression of
the commonly employed genus name Micrognathus Duncker (type-species,
Syngnathus brevirostris Ruppell) and the introduction of new names for the spec-
imens described as Corythoichthys brevirostris by Kaup (1856) and for the other
Species presently referred to Micrognathus. Until such time as a pipefish is found
which is clearly identifiable with Ruppell’s description, it seems best to consider
his description as poorly written and to treat specimens described by Kaup (1856)
as representative of Syngnathus brevirostris Ruppell. Kaup (1856) gives counts
of 14-15 + 30 rings, 5 subdorsal rings, 12 dorsal-fin rays (obviously in error), and
notes the presence of this species in the museums at Frankfurt, London, Stuttgart,

VOLUME 95, NUMBER 4 667

Fig. 4. Micrognathus brevirostris brevirostris. Top, Late yolk-sac larva (ca. 5.8 mm SL), GCRL
18155. Bottom, Planktonic juvenile (14.9 mm SL), GCRL 17428.

Berlin and Darmstadt. I am unaware of the fate of the specimens at Stuttgart,
Berlin and Darmstadt, but there are now 7 possible syntypes, collected by Ruppell
at Massaua and identified with S. brevirostris by Kaup, in the London and Frank-
furt collections. An adult male (BMNH 1860.11.9.60), accompanied by a hand-
written internal jar label stating “‘one of the typical specimens,’ has 19 dorsal-
_ fin rays and retains traces of patches of melanophores on or near the lateral,
inferior and median ventral ridges of the 8—9 anterior trunk rings (Fig. 3). The
Six remaining specimens, all distorted and in relatively poor condition, consist of
one male (SMF 902) and a lot of 5, including one male (SMF 4909-4913).

Among material examined, the brood pouch is developed below the 9-14 (* =
11.5) anterior tail rings in 19 males (33—51.5 mm SL), and the smallest brooding
male is 33 mm SL. Brood-pouch eggs are usually deposited in 1—2 layers and in
two transverse rows. A 36 mm male has 30 eggs in a 10-ring pouch, a 44 mm fish
has a single layer of 32 eggs through 12 of 13 pouch rings, and another (42 mm)
has 67 early embryos in a 2 X 2 arrangement in a 14-ring pouch. Although late
yolk-sac larvae are seldom found in preserved males of any species of Micro-
gnathus, 4-5 nearly straight larvae, ca. 5.8 mm SL (Fig. 4), were found with
several early (coiled) larvae in the partially filled pouch of a 41 mm fish. Config-
uration of principal body ridges is not distinguishable in these late larvae, but
the dorsal and caudal fins are well developed and the pectoral and anal fins are
present. Planktonic young (13-15 mm SL) lack dermal flaps, they have pointed
posterior angles on most rings, and the dorsal-fin base is a little elevated (Fig. 4).
Present materials show no evidence of geographic variation in coloration or me-
ristic values.

Many literature records of M. brevirostris from the Red Sea are based, wholly
or in part, on misidentifications of M. andersonii. Most have not been verified
here but two deserve comment. Four fish from Koseir (MCZ 3814, ZMB 7906),
collected by Klunzinger and apparently reported as M. brevirostris (Klunzinger
1871), are specimens of M. andersonii. Similarly, the Red Sea specimen of Dr.
Jousseaume (MNHN 93064), reported by Dollfus and Petit (1938), is also a mis-
identified example of M. andersonii.

Distribution.—Micrognathus b. brevirostris is a Red Sea endemic (Fig. 5) known

668 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

20° 40° 60° 80° 100° 120° 140° 160° 180° 160°
=

_J20°

=|209

b. pygmaeus
micronotopterus
atans

b. brevirostris
n

> M.
@ M.
* M.
6 M.

Fig. 5. Distributions of selected species of Micrognathus based on material examined. A record
of M. b. pygmaeus from Tahiti (ca. 17°32'S, 149°34’W) is omitted.

from the Gulfs of Suez and Aqaba in the north to the Strait of Perim in the south
(ca. 12°36’N, 43°16’E). Among present material, there is one lot of offshore plank-
tonic young and two lots of adults taken from clumps of coral (Stylophora sp.)
collected in 1.5 m. Data accompanying other samples lack useful information on
habitat or depth of capture. It is uncertain whether Fishelson’s (1971) reference
to the occurrence of M. brevirostris in the “‘Halophila .../Asymmetron .. .
community’’ (2—5 to ca. 40 m) is based on this species or its only Red Sea con-
gener, M. andersonii.

Material examined.—Forty-two specimens (excluding larvae) 13-62 mm SL,
including 7 possible syntypes*.

RED SEA, Gulf of Suez: ZMB 20003 (1, 48), ZMB 20004 (2, 44.5—45.5). Gulf
of Aqaba: HUJ F.2096 (1, 16.5), HUJ F.6522 (5,_31-36), HUJ F.9258 C1, 17),
TAU P.6265 (1, 26). Sinafur Is.: BMNH 1951.1.16.92—94 (3, 36.547). Al Ghar-
daqa (27°13’N, 33°51’E): GCRL 18154 (7, 41-52), GCRL 18155 (5, 41-62). Quseir:
NMW 40279 (1, 37.5). Jiddah: RMNH 28610 (1, 40.5). Farasan Is.: SMF 4907 (1,
38.5). Massaua: BMNH 1860.11.9.60 (1, 51.5*), SMF 902 (1, ca. 52.5*), SMF
4909-4913 (5, 45.5-53.5*). Ethiopia: GCRL 17428 (2, 13-15). Loc. uncertain:
MNHN 52-302 (4, 36.5—46.5), Calypso, no other data.

Micrognathus brevirostris pygmaeus Fritzsche
Figs. 6, 7

Micrognathus pygmaeus Fritzsche, 1981:771, fig. 1 (orig. descr.; Taone, Tahiti).

Diagnosis.—Subdorsal tail rings usually 2.75—3.75, pectoral-fin rays usually 11-
12, side of head typically without narrow pale bars, subadults and adults usually
with minute dark spots on opercle.

Description.—Rings 15—16 + 28-31 (15 + 28-31 in 99%), dorsal-fin rays 18-
21, pectoral-fin rays 10-13 (usually 11-12), anal-fin rays usually 3, subdorsal rings

VOLUME 95, NUMBER 4 669

Fig. 6. Micrognathus brevirostris pygmaeus. Lateral and dorsal aspects of head and anterior trunk
rings, together with section of body illustrating ridge configuration and dorsal and anal fins. From
46.5 mm SL brooding male (USNM 215313).

1.5-0.25 + 2.75—4.25 = 3.75-5.0 (see Tables 1-3 for additional counts). Propor-
tional data, based on 27 specimens 26.0—55.0 @ = 45.7) mm SL, follow: HL in
SL 7.2—10.2 (9.2), snout length in HL 2.6—3.3 (3.0), snout depth in snout length
1.8-3.0 (2.2), length of dorsal-fin base in HL 1.1-1.6 (1.3), pectoral-fin length in
HL 3.7—-5.9 (4.7), length of pectoral-fin base in pectoral-fin length 1.4—-2.0 (1.7),
anal ring depth in HL 2.2-3.3 (2.6), trunk depth in HL 1.9-2.8 (2.3). Distal half
or more of tail with principal ridges flared laterad and with posterior angles of
rings produced to form hook-like points (Fig. Ic); dermal flaps typically short
(Fig. 6), often with frilled margins or with short distal branches.

Ground color pale to dark brown. Pale fish often with a brown lateral stripe on

670 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 7. Micrognathus brevirostris pygmaeus. Top and middle, USNM 217478 (42 mm SL, adult
female). Bottom, AMS 1.20770-004 (52 mm SL, male).

posterior half of snout which continues on postorbital and expands to encompass
most or all of opercle; median portion of opercle usually with tan ocellus-like
blotch, often narrowly margined with pale, and (in specimens >30 mm SL) usually
with 1-20 discrete, minute, brown to near-black spots (Fig. 7); trunk and tail
plain, with indications of narrow dusky bars on lower part of side and venter of
trunk, or with similar bars encircling trunk; fins mainly hyaline. Dark specimens
often with a narrow, transverse, pale bar below angle of gape; eye usually with
dark bars radiating from pupil; sometimes with two pale bars below orbit; opercle
much as described for pale fish; body usually with 10-11 irregular pale bars
crossing dorsum and upper part of side; sometimes with near-black blotches on
lateral ridge of 3rd—4th and 7th—8th trunk rings; lower part of side of trunk and
venter of trunk and tail with well-defined or diffuse pale bars; brood-pouch folds
may have narrow pale bars extending ventrad from middle of each pouch ring
and narrow mottled edging along the free margins; fins usually flecked or shaded
with brown.

VOLUME 95, NUMBER 4 671

Comparisons.—See this section under M. b. brevirostris.

Remarks.—Ogilby (1908) described Corythoichthys spinicaudatus from a single
brooding male (57 mm TL) taken at Cape York in southern Queensland. He stated
that the holotype was in the Queensland State Museum, but it was evidently
never cataloged therein, it cannot now be located in any collection, and is ap-
parently lost (R. J. McKay, pers. comm.). This specimen was described as having
‘fall the caudal ridges strongly spinigerous throughout their length,’ 16 + 30
rings, 22 dorsal-fin rays, 14 pectoral-fin rays, 14 brood-pouch rings, and ‘‘dorsal
fin inserted on 2 body and 3 caudal rings.’’ The description is neither illustrated
nor diagnostic and the identity of the holotype is uncertain. The population de-
scribed here agrees with Ogilby’s holotype in having ‘‘spinigerous’’ tail ridges
and occurs in northern Queensland. On the other hand, Ogilby’s meristic data
best agree with those of Micrognathus andersonii, but specimens of this species
seldom have tail ridges which could be termed “‘spinigerous’’ and M. andersonii
is not known to occur near Cape York. Although Duncker (1915) and subsequent
authors have referred Ogilby’s name to the synonymy of M. brevirostris or listed
it, without comment, as a subspecies thereof (Whitley and Allan 1958), Cory-
thoichthys spinicaudatus 1s a nomen dubium.

Fritzsche (1981) described Micrognathus pygmaeus from two apparently faded
or bleached specimens, including a brooding male (23 mm SL, holotype) and a
presumed female paratype (20 mm SL), collected at Tahiti. The species was
_ distinguished from M. brevirostris on the basis of fewer trunk rings (14 versus
15-17), a lower HL in SL ratio (7.1-7.4 versus 8.0-10.2), a higher snout length
in HL ratio (3.2—3.4 versus 2.2—3.2) and its small size at maturity. I have examined
these fish and find that both have 15 rather than 14 trunk rings and that the
principal tail ridges are flared laterad (Fig. Ic). Although both specimens were
described as having | + 4 subdorsal rings, the anterodorsal margin of the Ist tail
ring 1s curved atypically anteriad in the paratype so that the subdorsal rings are
0 + 4 in dorsal aspect or 0.25 + 4 when viewed from the side. Since proportional
and meristic values, as well as other morphological features, do not differ signif-
icantly from the population described here, I consider pygmaeus to be the first
available name for the western Pacific subspecies of M. brevirostris.

The small size of the mature holotype is noteworthy, but two other small adult
males are included in examined material. One (25 mm SL) from Bikini Atoll
(USNM 141197), reported as M. brevirostris by Herald (1953), has pouch eggs
through 10 of 12 pouch rings and retains traces of bars on the lower part of the
trunk. The other (22 mm SL) is from the vicinity of Raine Is. (11°36’N, 144°01’E),
N Queensland (AMS I.20757-001). This specimen has ca. 10 pouch eggs through
10 of 11 pouch rings, apparently 9 x 10 pectoral-fin rays (omitted from Tables),
and there are dark spots, indicative of bars, on the lateral and inferior trunk
ridges. Except for these specimens, the smallest male with evidence of a devel-
oping brood pouch and the smallest brooding male are both 39.5 mm SL. The
pouch extends below 11-14 @& = 12.2) rings in 22 brooding males (39.5—55 mm
SL), and the brood-pouch eggs are usually in a single layer of 2-4 transverse
rows. A 49.5 mm SL fish has a total of 37 eggs distributed through the 10 anterior
rings of a 12-ring pouch.

Dark spots are absent from the opercle of most small fish, but they are usually
present on specimens larger than 30 mm SL and spots are usually more numerous
on mature males. Among recently preserved fish (41-50 mm SL), there are 6-20

672 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(x = 12.8) spots on each opercle of 18 males and 0-12 (x = 4.8) on 23 presumed
females.

Distribution.—Micrognathus b. pygmaeus is known from eastern Indonesia
and northern Queensland to Bikini Atoll and Tahiti (Fig. 5). Occurrence at New
Caledonia is based on the provisional identification of two bleached specimens.
Available materials indicate that M. b. pygmaeus is replaced by M. andersonii
in southern Queensland and that these species are sympatric within the region
from Cape Melville (ca. 14°10’S) to the Endeavour Reef area (ca. 15°47'S). These
species have occurred together in samples from Indonesia, as well as from Two
Isles (ca. 15°01'S)-and Little Hope Is. (ca. 15°47’'S), Queensland. Among collec-
tions with adequate data, two are from “‘reef pools,” five from 0-1.1 m, and
seven from I—8 m; maximum range for any collection is 0—20 m.

Material examined.—Seventy-eight specimens, 19-55 mm SL, including Ho-
lotype (USNM 207933, 23 mm SL, brooding male, Society Is., Tahiti, Taone,
1965), and Paratype (USNM 215775, 20, presumed female, data as for holotype).
INDONESIA, Buru Is.: USNM 94084 (1, 45.5, paratype of Syngnathus microno-
topterus). Ceram: USNM 215313 (1, 46.5). Banda Is.: ZMA 116.478 (1, 35). Irian
Jaya: RMNH 27534 (1, 34). PAPUA NEW GUINEA: USNM 222933 (1, 39.5).
PALAU IS.: CAS 47888 (1, 46), CAS 47891 (1, 49), CAS 47892 (1, 40.5), CAS
47896 (1, 47), CAS 47898 (1, 28.5), CAS 47903 (1, 47), CAS 47904 (1, 44). AUS-
TRALIA, Queensland: AMNH 35895 (1, 42), AMNH 35896 (3, 40-48), AMNH
35899 (4, 28-47.5), AMNH 35900 (3, 49-52), AMNH 35902 (1, 53), AMNH 35904
(4, 23-49), AMNH 35915 (1, 20.5), AMS 1.14030 (1, 45.5), AMS I.18755-001 (1,
49.5), AMS I.19444-034 (2, 41-55), AMS I.19461-014 (4, 45.5—46), AMS I.19473-
049 (3, 19-50), AMS I.19473-183 (1, 26), AMS I.20756-001 (1, 19), AMS 1I.20757-
001 (1, 22), AMS I.20770-004 (4, 47-52), AMS I.20782-001 (1, 53), AMS I.21422-
066 (1, 48.5), AMS I.21539-067 (1, 47.5), ANSP 119351 (3, 38.5—46), ANSP 148774
(2, 49-50.5), GCRL 17879 (5, 42-47), SIO 61-132 (2, 48-52). BISMARCK ARCH. :
USNM 217478 (1, 42). SOLOMON IS.: CAS 19951 (1, 32.5), CAS-SU 25155 (1,
39.5), GCRL 13780 (1, 46), USNM 214140 (4, 39.5—45). SANTA CRUZ IS.: CAS
47899 (1, 39.5). NEW HEBRIDES IS.: CAS 19952 (1, 30.5). NEW CALEDONIA:
USNM 215307 (2, 3440.5). MARSHALL IS., Bikini Atoll: USNM 141197 (2,
25—25.7).

Micrognathus andersonii (Bleeker)
Figs. 8, 9

Syngnathus Andersonii Bleeker, 1858:465 [orig. descr.; Nova-selma, Kokos (Co-
cos) Is. (Indonesia)].

Corythroichthys tanakae Jordan and Starks, 1906:696, fig. 2 (orig. descr.; Tane-
gashima, Japan).

Diagnosis.—Trunk rings modally 16, principal ridges not angled laterad on
distal half of tail, dorsal-fin rays usually 18—22, subdorsal tail rings usually 2.75—
4.5, snout length averages 2.9 in HL, anal ring depth averages 2.4 in HL, often
with prominent dark blotches above lateral trunk ridge.

Description.—Rings 15—17 + 27-32 (16-17 + 28-32 in 99%), dorsal-fin rays 17—
24, pectoral-fin rays 11-13, anal-fin rays usually 3, subdorsal rings 1.75—0.25 +

VOLUME 95, NUMBER 4

Fig. 8. Micrognathus andersonii. Lateral and dorsal aspects of head and anterior trunk rings,

together with section of body illustrating ridge configuration and dorsal and anal fins. From 57 mm
SL brooding male (HUJ F.8034).

2.5—5.0 = 3.75—5.75 (see Tables 1-3 for additional counts). Proportional data,
based on 56 specimens 37.5—75.0 (x = 53.2) mm SL, follow: HL in SL 8.1-10.4
(9.0), snout length in HL 2.7-3.4 (2.9), snout depth in snout length 1.6—2.8 (2.1),
length of dorsal-fin base in HL 0.9-1.5 (1.2), pectoral-fin length in HL 2.8—5.3
(4.0), length of pectoral-fin base in pectoral-fin length 1.3—2.4 (1.7), anal ring depth
in HL 1.9-3.1 (2.4), trunk depth in HL 1.7-2.8 (2.0). Principal tail ridges essen-
tially straight (Fig. la), infrequently elevated slightly and with pointed posterior
angles (Fig. Ib) but not clearly angled or flared laterad with enlarged hook-like

674 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig.9. Upper pair, Micrognathus andersonii, GCRL 17020: Top, 64mm SL, male, pale coloration;
Bottom, 72 mm SL, male, dark coloration. Lower pair, M. micronotopterus: Top, USNM 94082 (55
mm SL, brooding male, holotype); Bottom, CAS-SU 30916 (55 mm SL, male).

posterior angles (Fig. Ic). Dermal flaps usually short (Fig. 8), simple, with frilled
margins or with short branches distally.

Ground color pale to dark brown (Fig. 9); variously plain, mottled or blotched
and spotted with pale or brown; eye usually with dark bars radiating from pupil;
often with 1-3 pale or brown bars below eye; opercle usually plain, sometimes
with a pale blotch or bar but without discrete dark spots; body often with 9-13

VOLUME 95, NUMBER 4 675

narrow bars crossing dorsum and part of side from nape to caudal-fin base; with
or without 1-4 dark blotches between lateral and superior trunk ridges; typically
without bars crossing lower half of trunk; dorsal and pectoral fins hyaline or with
fin-rays and membrane streaked or flecked with brown; caudal fin usually brown-
ish with pale distal margin.

Comparisons.—The combination of modally 16 trunk rings, usually 18—22 dor-
sal-fin rays, short snout and essentially straight tail ridges distinguishes M. an-
dersonii from congeners. In addition, adult males of M. andersonii tend to have
more brood-pouch rings than males of Indo-Pacific congeners (13-18 versus 10-
15). Infrequent specimens with atypical counts of 15 trunk rings are distinguished
from M. micronotopterus and M. natans, species sharing a count of 15 trunk
rings and essentially straight tail ridges, by a lower average snout depth in snout
length ratio (2.1 versus 2.6—3.4), a lower average length of dorsal-fin base in HL
ratio (1.2 versus 1.6—1.7) and other features (see descriptions). Specimens of M.
andersonii and M. micronotopterus may have prominent dark blotches above the
lateral trunk ridge and these sympatric species cannot be distinguished solely on
the basis of preserved coloration. Small specimens of M. andersonii (ca. 12-20
mm SL) are best distinguished from those of Indo-Pacific congeners by their
typically higher frequency of trunk rings (usually 16-17 versus 14—15) and other
meristic differences (see Tables 1-3).

Remarks.—Although clearly representing a separate taxon, this species was
_ referred to the synonymy of M. brevirostris by Weber (1913) and it has been so
treated by subsequent authors.

Bleeker’s (1858) unillustrated original description of Syngnathus andersonii,
based on a single male specimen, gives counts of 16 + 27 or 28 rings, | + 4
subdorsal rings, 19 or 20 dorsal-fin rays, 14 pectoral-fin rays, 3 or 4 anal-fin rays,
10 caudal-fin rays and 14 or 15 pouch rings. The holotype (RMNH 7227), now 47
mm SL, has 20 or 21 rays in the damaged dorsal fin, 10 caudal-fin rays, 16 + 28
rings, 0.75 + 4.25 subdorsal rings, 15 pouch rings and some dermal flaps persist
on the head. Both pectoral fins are damaged but Bleeker’s count of 14 fin-rays is
probably in error, since the number is 11-13 in fins examined here (Table 2).
Aspects of the history of the holotype were discussed by Dawson (1977) but there
is little doubt that this specimen, now faded and pencil-marked, was the model
for Bleeker’s unpublished Atlas illustration of this species (Pl. 450, fig. 3).

Syngnathus sundaicus, described from Java by Bleeker (1853) and referred to
the synonymy of Micrognathus brevirostris by Duncker (1915), may have been
based on an Indonesian species of Micrognathus (andersonii, micronotopterus,
or b. pygmaeus) but the name must be considered a nomen dubium. The unfigured
original description, based on a van Hasselt drawing which has since been lost,
is not diagnostic and the name cannot be referred with certainty to any genus or
species of pipefish.

Jordan and Starks (1906) based their description of Corythroichthys tanakae
on eight specimens (including 5 males) and stated that the “‘type’’ was numbered
USNM 53271 and that a ‘“‘cotype’’ was numbered SU 9358. Springer and Esch-
meyer (1974) have shown that these numbers were apparently transposed and
that the holotype is presently numbered CAS-SU 9358. This species was referred
to the synonymy of Micrognathus brevirostris by Duncker (1915) and it has been
so treated by most subsequent authors. Although I have no information on the

676 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

fate of four of the eight original specimens, I find the holotype and three presumed
paratypes (USNM 53271), including two males and a female, to be conspecific
with M. andersonii.

The smallest examined male with evidence of a developing brood pouch is 33.5
mm SL and the smallest brooding fish is 34 mm SL. The brood pouch extends
below 13-18 (* = 15.4) tail rings in 70 brooding males (34—70.5 mm SL) and the
brood-pouch eggs are usually deposited in one layer of 2—7 transverse rows. A
58.5 mm male has a total of 95 eggs deposited through the anterior 13 rings of a
14-ring pouch, and a 67.5 mm fish has 103 eggs through 14 of 15 pouch rings.
Dermal flaps are usually short (Fig. 8) and often have frilled margins or short
distal branches. Infrequent samples include occasional subadult-adults with lon-
ger flaps which are somewhat similar to those of M. micronotopterus. Preserved
coloration is highly variable and both pale and dark fish, without regard to size
or sex, may occur in the same collection. Some young fish (15.5—16 mm SL) from
inshore plankton collections (GCRL 16873, 17452) have dermal flaps on the body
and the adult pattern of bars on the dorsum.

Present materials show little geographic variation in meristic values but some
differences are noteworthy. Numbers of tail rings are lowest (27-30) in fish from
the Philippines and the Palau Is. and highest (29-32) in specimens from Australia
and the Ryukyu Is. Similarly, Philippine fish have the lowest numbers of dorsal-
fin rays (17-21) and highest numbers (19-24) occur in specimens from the Ryukyu
Is. Pectoral-fin rays are most often I1 (58% of 142 counts) in fish from the Red
Sea and western Indian Ocean but usually 12-13 (92% of 388) in other material.
In addition to having low numbers of tail rings and dorsal-fin rays, examined
Philippine specimens do not exceed 45 mm SL and brooding males are smaller
than those from other areas (34—45 versus 41-70 mm or longer). Despite these
differences, I find no basis for separate taxonomic treatment of the Philippine
population.

Two fish with atypical meristic values (not in Tables) are provisionally referred
to M. andersonii. A bleached female (65 mm SL) from New Caledonia with
somewhat flared tail ridges and without persistent dermal flaps seems to best
agree with this species, but there are 14 rays in each pectoral fin, 17 dorsal-fin
rays and 15 + 29 rings. The condition of the specimen, coupled with high numbers
of pectoral-fin rays and low numbers of trunk rings and dorsal-fin rays prevents
positive identification. The second questionable fish is an early juvenile (ca. 12
mm SL) taken with a specimen of M. andersonii (14 mm SL) by the ALBATROSS
at Ticao Is., Philippine Is. (USNM 133054). Presence of 15 + 29 rings would
indicate that this is an example of M. micronotopterus, but counts of 20 dorsal-
fin rays and 1.25 + 3.75 subdorsal rings are high for that species (see Tables I-
3) and suggest that this specimen is best referred to M. andersonii.

Distribution.—Micrognathus andersonii is known from the Red Sea (Gulfs of
Suez and Aqaba to the Dahlak Arch.) and western Indian Ocean to Japan (Honshu
Is.) and southeastward to Samoa and the Tonga Is. (Fig. 10). Examined Indian
Ocean material confirms the occurrence of this species on the African coast from
Kenya to South Africa (ca. 32°10’S) and there are specimens from Madagascar,
the southeastern tip of India and western Indonesia. There are no specimens of
any species of Micrognathus in numerous collections examined from Aldabra and
the Comoro Is., the Seychelles and Amirante Is., St. Brandon (Cargados Carajos)

VOLUME 95, NUMBER 4 677

20° 40° 60° 80° 100° 120° 140° 160° 180° 160°
Sees ay ey

M. andersonii

Fig. 10. Distribution of Micrognathus andersonii based on material examined.

Shoal, Mauritius, the Maldive Is., Chagos Archipelago, Sri Lanka, and the Cocos-
Keeling Is. Records of M. brevirostris from Aldabra and the Seychelles (Smith
1955, 1963; Smith and Smith 1963) are not supported by material in collections,
and are, at least in part, based on misidentified specimens of a superficially similar
pipefish, Phoxocampus belcheri (Kaup). A listing by Marshall (1950) is based on
the misidentification of the type-locality as Cocos-Keeling rather than the Cocos
Is. of Indonesia. These observations indicate the absence or a low level of abund-
ance of M. andersonii and other species of Micrognathus at these Indian Ocean
localities. In Australia, M. andersonii is known only from the vicinity of Cape
Melville (ca. 14°12’S) and southward along the Queensland coast to the vicinity
of Southport (ca. 27°58’S). This species is sympatric with M. b. brevirostris in
the Red Sea, with M. micronotopterus in Indonesia and the Philippine Is., and
with M. b. pygmaeus on the Queensland coast of Australia (ca. 14—16°S), as well
as at a number of other western Pacific localities from Indonesia to Palau and
southeastward to the New Hebrides and New Caledonia. Thirty samples with
useful data indicate collections of juveniles-adults in rock or reef tidepools, among
‘‘turtle grass’’ and seaweed, and on reef and sand flats. Maximum recorded depth
range is 0-5.2 m, but 29 of these samples are from 0-2 m. Planktonic young have
been taken in the upper 0-100 m over depths to 4825 m.

Material examined.—Five hundred eighty-three specimens, 12-78 mm SL, in-
cluding the holotype (RMNH 7227, 47.0, male, Nova-selma, Kokos (Cocos) Is.,
Indonesia). RED SEA, Gulf of Suez: BMNH 1925.9.19.35 (2, 39-54.5), BMNH
1925.9.19.36 (1, 56.5), BMNH 1925.12.31.21 (1, 65.5), HUJ F.2104 (1, 65), HUJ
F.8034 (3, 49-62), HUJ F.9243 (1, 59.5), TAU P.5653 (3, 51.5—63.5), ZMB 31293
(1, 54). Gulf of Aqaba: BMNH 1960.3.15.99 (1, 50.5), GCRL 13880 (3, 54.5—57),
HUJ F.4748 (3, 48—-56.5), HUJ F.8032 (2, 50.5—58.5), HUJ F.8033 (18, 4459.5),
HUJ F.9244 (1, 39.5), TAU P.3560 (1, 35), TAU P.6264 (1, 39.5), TAU P.6266
(1, 32.5). Quseir (Koseir): MCZ 3814 (3, 54.5-55), ZMB 7906 (1, 55). Suakin:
BMNH 1886.6.9.27 (7, 24.5—65.5). Dahlak Arch.: HUJ F.9245 (2, 33.535). Jid-

678 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

dah: RMNH 18391 (2, 58.5-63.5), RMNH 18392 (2, 43-48.5). Loc. uncertain:
HUJ F.8028 (1, 59.5), HUJ F.8031 (1, 54), HUJ F.9246 (1, 21.5), MNHN 93-64
(1, 52.5), TAU P.3557 (2, 36.5—5S6.5). INDIAN OCEAN, Kenya: RUSI 4664 (1,
27.5), RUSI 15029 (1, 41), RUSI 15037 (3, 14-60.5), RUSI 15040 (1, 44). Zanzibar:
MCZ 36592 (1, 48), RUSI 4109 (1, 47), RUSI 15030 (7, 43.5-62). Mozambique:
RUSI 4100 (1, 57.5), RUSI 4108 (1, 33.5), RUSI 12241 (1, 70.5), RUSI 12242 (1,
50.5), RUSI 12244 (1, 45), RUSI 15031 (1, 46.5), RUSI 15032 (1, 51.5), RUSI
15033 (3, 44-64), RUSI 15034 (4, 41-57), RUSI 15035 (12, 35-67.5), RUSI 15036
(2, 57.5—61), RUSI 15041 (3, 52-64), RUSI 15042 (6, 33-61), SAM 28989 (1, 36.5).
South Africa: RUSI 15038 (30, 32.5—65), RUSI 15043 (10, 15-59). Madagascar,
Nosi Bé: MNHN 31:263 (1, 43.5), USNM 214141 (1, 38). India, Musaltiva Is.:
FMNH 75853 (1, 46.5). Loc. uncertain: MNHN 4956 (1, 63), RUSI 12275 (9,
32.5—50.5). PACIFIC OCEAN, Indonesia: BMNH 1867.11.28.355 (1, 58), BMNH
1880.7.12.7-10 (4, 46—-52.5), RMNH 21106 (1, 37), USNM 94083 (1, 48.5) and
USNM 94086 (1, 43), paratypes of Syngnathus micronotopterus, USNM 210654
(1, 52), USNM 215287 (1, 36), ZMA 115.977 (1, 43.5), ZMA 115.978 (2, 38.544),
ZMA 115.979 (1, 22), ZMA 115.981 (1, 71.5), ZMA 116.039 (2, 31.5-—35), ZMUC
P.39598 (1, 16.5). Hong Kong: CAS 47902 (2, 44-49). Philippine Is.: CAS 47886
(1, 33.5), CAS-SU 29952 (1, 33.5), CAS-SU 39164 (1, 40), CAS-SU 51498 (5, 36—
45), GCRL 16999 (1, 15.6), USNM 94087 (40, paratype of S. micronotopterus),
USNM 133054 (2, 12-14), USNM 219514 (23, 24.5S—42.5), USNM 220656 (1, 32),
UWCE 20835 (1, 39.5), UWCF 20836 (1, 36.5), ZMUC P.39596 (1, 15), ZMUC
P.39597 (1, 12.5). Ryukyu Is.: CAS-SU 9358 (68, male, holotype of Corythro-
ichthys tanakae), CAS-SU 9782 (13, 35—75), FMNH 42866 (1, 62.5), USNM 53271
(3, 56-60, presumed paratypes of C. tanakae), USNM 70741 (78, 31.5—72), USNM
172063 (1, 60.5), YCM P.1471 @Q, 28.5-57), YCM P.1765 (1, 51), YCM P.2859 (1,
33.5), YCM P.6007 (1, 28.5), YCM P.6010 (4, 40-52.5), YCM uncat. (1, 63). Japan,
Honshu Is.: YCM P.7981 (1, 71). Palau Is.: BPBM 9596 (4, 48-60.5), CAS 34089
(1, 49.5), CAS 47887 (1, 32.5), CAS 47889 (2, 28-28.5), CAS 47890 (1, 40.5), CAS
47893 (1, 45.5), CAS 47894 (3, 38-45), CAS 47895 (1, 28), CAS 47897 (1, 51.5),
CAS 47905 (1, 24). Mariana Is.: AMNH 27006 (1, 27), ANSP 80661 (1, 45), ANSP
90783 (1, 49.5), CAS 34088 (1, 22.5), CAS 47885 (1, 22.5), CAS 47900 (1, 43.5),
CAS 47901 (1, 41), FMNH 42864 (1, 24), FMNH 42865 (1, 51.5), UG 1556 (1,
43), UG 6260 (1, 45.5), UMMZ 197958 (1, 40.5), UMMZ 197992 (1, 31.5), UMMZ
198004 (2, 49-53.5), UMMZ 198224 (1, 42), USNM 109382 (1, damaged), USNM
154631 (6, 40-47.5). Papua New Guinea: KFRS F.2956 (1, 36). Australia, Queens-
land: AMNH 35897 (1, 51), AMNH 35903 (7, 33.5-—65), AMNH 43098 (1, 65),
AMS 1.2962 (4, 30-55.5), AMS 1.6214-5 (2, 57-63.5), AMS 1.11429 (5, 50.5—57),
AMS I.12972 (1, 55.5), AMS I.18784-001 (3, 57.5—58), AMS I.19477-066 (1, 42.5),
AMS 1.20200-021 (3, 54-67), AMS 1I.20207-010 (2, 47.5—S1.5), AMS 1.20463-041
(2, 60.5-61), AMS 20760-001 (1, 75), AMS 1.20761-001 (1, 53.5), AMS I.20772-
003 (3, 54-61), AMS I.20773-006 (1, 38.5), AMS I.20780-004 (7, 46-62.5), AMS
1.20787-006 (1, 33.5), AMS 1.22851 (2, 40.5—43), AMS 1I.21538-022 (1, 55), AMS
[A.229 (7, 46.5-61.5), AMS IA.1807 (1, 58.5), AMS IA.2624 (8, 22.5—60), AMS
[A.4821 (1, 70.5), AMS IA.4822 (1, 78), AMS IA.5645 (1, 49), AMS IA.7066 (1,
65), AMS I[A.7364-65 (11, 45.5—69.5), AMS IB.6580 (2, 54-56), AMS IB.6686 (1,
57.5), ANSP 113480 (9, 50-72.5), BMNH 1873.4.3.76-77 (1, damaged), BMNH
1933.1.25.11 (1, 70), BMNH 1933.1.25.12-13 (2, 56-58), BPBM 14344 (1, 39), CAS

VOLUME 95, NUMBER 4 679

13784 (4, 62.5—64), CAS 13789 (1, 67.5), CSIRO B.1366 (6, 50.5—58), GCRL 15533
(1, 59.5), GCRL 16122 (1, 68), GCRL 16300 (2, 51-56), GCRL 16873 (1, 16),
GCRL 17020 (5, 45-72.5), GCRL 17451 (1, 43), GCRL 17452 (1, 15.5), MCZ 36896
(1, 58), QM 1.2050 (1, 50), QM 1.6585 (1, 53), QM 1.9318 (11, 35-68), QM 1.9381
(7, 41-56), SIO 61-132 (7, 48-56), USNM 177141 (1, 47.5), USNM 231699 (5,
51.5-61), USNM 231700 (2, 51.5—56.5), USNM 231701 (2, 44-53), WAM P.26499-
001 (1, 41.5). Ponape: USNM 223108 (1, 37). Solomon Is.: BPBM 15678 (1, 36.5),
CAS 20025 (1, 44). New Hebrides: AMS 1.6364 (1, 36), AMS IA.772 (1, 32.5),
GCRL 13781 (2, 46.5—50), USNM 214139 (3, 3441). New Caledonia: AMS I.18446-
001 (1, 49), CAS 19946 (3, 36-39), CAS 19953 (14, 42-58), CAS 19954 (2, 49-52),
CAS 20027 (3, 45-55.5), MNHN 9297 (1, 65). Gilbert Is. (Tarawa): AMS I.18054-
008 (1, 51). Fiji Is.: CAS-SU 24863 (1, 43). Samoa Is.: GCRL 16290 (2, 37.5—46).
Tonga Is.: USNM 83004 (1, damaged). Loc. uncertain: MNHN A.477 (1, 65),
‘‘Poulo Condor’’ (Indochina).

Micrognathus micronotopterus (Fowler)
Figs. 9, 11

Synenathus micronotopterus Fowler, 1938:42, fig. 14 (in part, orig. descr.; Can-
imo Is., S Luzon, Philippine Is.).

Diagnosis.—Trunk rings modally 15, principal ridges not angled laterad on
distal half of tail, dorsal-fin rays usually 17-18, subdorsal tail rings usually 2.25—
3.5, snout length averages 2.6 in HL, anal ring depth averages 2.8 in HL, often
with prominent dark blotches above lateral trunk ridge.

Description.—Rings 15 + 28-31, dorsal-fin rays 17-19, pectoral-fin rays 10—13
(usually 11-12), anal-fin rays usually 3, subdorsal rings 1.5—0.25 + 2.25-3.75 =
3.25—4.5 (see Tables 1-3 for additional counts). Proportional data, based on 19
specimens 28.0-57.0 (« = 47.3) mm SL, follow: HL in SL 7.2-8.9 (8.3), snout
length in HL 2.3-3.0 (2.6), snout depth in snout length 2.1—3.1 (2.6), length of
dorsal-fin base in HL 1.3—1.9 (1.6), pectoral-fin length in HL 3.9-5.6 (4.9), length
of pectoral-fin base in pectoral-fin length 1.4—2.1 (1.7), anal ring depth in HL 2.5-
3.2 (2.8), trunk depth in HL 2.1-2.9 (2.4). Principal tail ridges essentially straight
(Fig. la), infrequently elevated slightly (Fig. 1b), not angled or flared laterad with
posterior angles of rings produced to prominent hook-like points. Dermal flaps
on head of adults mostly long, slender and unbranched (Fig. 11).

Ground color tan to dark brown, usually mottled; usually with 10-12 narrow,
diffuse, pale bars crossing dorsum between nape and caudal-fin base; sometimes
with indications of subequal dark and pale bars on lower half of trunk; tan to
light brown fish usually with 1-6 diffuse dark blotches between lateral and su-
perior trunk ridges; dorsal fin often plain, sometimes with 2-3 irregular brownish
stripes crossing fin-rays and membrane; pectoral fin often barred or shaded with
brown; caudal fin brownish with pale distal margin.

Comparisons.—The combination of modally 15 trunk rings, usually 17—18 dor-
sal-fin rays, relatively long snout and essentially straight tail ridges distinguishes
subadult-adult specimens of M. micronotopterus from congeners. Furthermore,
the dermal flaps on the dorsum of head of adults (Fig. 11) are typically long and
simple, whereas those of adult congeners are typically shorter and often frilled
or with short branches distally. Occasional adult specimens of M. andersonii or

680 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 11. Micrognathus micronotopterus. Lateral and dorsal aspects of head and anterior trunk
rings, together with section of body illustrating ridge configuration and dorsal and anal fins. From 55
mm SL male (CAS-SU 30916).

M. natans with atypical counts of 15 trunk rings could be confused with M.
micronotopterus, but these can usually be distinguished without undue difficulty.
The HL in SL ratio of M. micronotopterus averages somewhat less than that of
M. andersonii (8.3 versus 9.0), the snout is more slender (snout depth in snout
length averages 2.6 versus 2.1) and specimens of M. andersonii seldom have long
head flaps (usually present in adult micronotopterus). In areas of sympatry, small
specimens of M. micronotopterus (< ca. 20 mm SL) are best distinguished from
juvenile M. andersonii by the number of trunk rings (modally, 15 versus 16 in
andersonii). Compared to M. natans, specimens of M. micronotopterus have a
lower average snout depth in snout length ratio (2.6 versus 3.4), a deeper body
(anal ring depth in HL averages 2.8 versus 4.8), fewer subdorsal tail rings (2.25—

VOLUME 95, NUMBER 4 681

3.75 versus 3.5—5.0) and often have dark blotches on the side of the trunk (absent
in natans).

Remarks.—Fowler’s (1938) description was based on the holotype (USNM
94082) from the Philippine Is. and ‘‘a series of paratypes from the Philippines
and the East Indies.’’ Of the six fish now catalogued as paratypes of M. micro-
notopterus, two (USNM 94085, 99007) are conspecific with the holotype, one
(USNM 94084) is a specimen of M. b. pygmaeus, and three (USNM 94083, 94086,
94087) are representatives of M. andersonii. The holotype (Fig. 9) is a brooding
male with 11 pouch rings, long simple flaps on the supraorbital, frontal and nuchal
ridges, persistent lateral blotches on the 4th, 9th and 13th trunk rings, and the
following measurements (mm): SL 55.5, HL 6.7, snout length 2.7, snout depth
0.9, length of dorsal-fin base 3.8, pectoral-fin length 1.3, length of pectoral-fin
base 0.6, anal ring depth 2.6, trunk depth 3.0 (see Tables 1-3 for additional data).
Herald and Randall (1972) identified ‘‘the two type specimens’’ of Ichthyocampus
annulatus Macleay as Micrognathus brevirostris and suggested that the figure
(Macleay, 1878, pl. 10, fig. 6) accompanying the original description (=Yozia
bicoarctata) was published in error. The two fish mentioned by Herald and Ran-
dall (AMS I.16288-001, formerly Macleay Mus. F.262) and a third ‘‘syntype”’
(AMS JA.1556) are conspecific with Micrognathus micronotopterus. 1 find no
direct evidence that Macleay identified these specimens as [chthyocampus an-
nulatus, and the original description (Macleay, 1878) does not fully agree with

‘these “‘syntypes’’ or with the figured specimen. However, except for numbers of
rings and dorsal-fin rays, the description agrees closely with the figure and with
the data from some 50 specimens of Yozia bicoarctata. In cases where a figured
description differs strikingly from the presumed type material, I believe that the
name must be applied to the illustrated taxon. Therefore, I consider Ichthyocam-
pus annulatus a junior synonym of Yozia bicoarctata (Bleeker) rather than a
senior synonym of Micrognathus micronotopterus.

The smallest examined male with evidence of a developing brood pouch is 28
mm SL, and the smallest brooding fish is 31.5 mm SL. The brood pouch is
developed below 11-15 (« = 12.7) tail rings in 10 brooding males (42-56 mm SL),
and the brood-pouch eggs are usually deposited in one layer of 24 transverse
rows. A 47 mm SL male has a total of 16 eggs in two rows through 8 rings of an
1l-ring pouch, whereas a 56 mm SL fish has ca. 92 eggs deposited in 4 rows
through 11 of the 13 pouch rings. Specimens faded in preservative, or those with
very dark ground color, may lack dark blotches above the lateral trunk ridge, but
others may have |-6 blotches on each side of the trunk; sixteen examined spec-
imens have three lateral blotches located on the 4th—Sth, 8th-10th, and 12th—14th
trunk rings.

Two small fish, 20.5—23 mm SL (USNM 231698), provisionally referred to M.
micronotopterus, have 15 trunk rings, 4.25—4.5 subdorsal rings, and were taken
with 23 specimens of M. andersonii (24.5—42.5 mm SL).

Distribution.—Micrognathus micronotopterus is known from Singapore, west-
ern Indonesia, the Philippine Is. and Australia (Fig. 5); a specimen labelled
‘‘Cochinchina’’ (MNHN 84-897) is of uncertain origin. In Australia, this is the
only species of Micrognathus known to occur on the northwestern coast from
the Exmouth Gulf, Western Australia (ca. 114°22’E) to the vicinity of Yirrkala

682 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Mission, Northern Territory (ca. 136°53'E). Data from 8 samples indicate occur-
rence in tidepool and coral reef habitats to depths of 1.5—5.5 m. This species is
sympatric with M. andersonii in Indonesia and the Philippine Is., but it does not
seem to occur with M. b. pygmaeus in either Indonesia or Australia.

Material examined.—Thirty-eight specimens, 20.5-57 mm SL, including ho-
lotype and two paratypes.

Holotype (USNM 94082, 55.5 mm, brooding male, Philippine Is., SE Luzon
Is. [W entrance to San Miguel Bay], Canimo Is., Daet Pt., 15 June 1909, ALBA-
TROSS). Paratypes (USNM 94085, Philippine Is.: 1, 31.5, male, Cebu Is., reef
opposite Cebu, 7 Apr. 1908, ALBATROSS; USNM 99007, 1, 51.5, female, taken
with holotype). SINGAPORE: BMNH 1952.12.16.11 (1, 47), CAS-SU 30916 (8,
44.5—57), FMNH 46923 (1, 56), GCRL 17945 (1, 55). INDONESIA, Java: RMNH
27592 (1, 47.5), UMMZ 183301 (1, 47), USNM 215316 (1, 53). Brunei: RMNH
12424 (2, 53-56). PHILIPPINE IS., Negros Is.: USNM 231698 (2, 20.5—23). AUS-
TRALIA, Western Australia: AMS I.17060-047 (4, 36-38), GCRL 18310 (1, 35.5),
QM 1.16658 (1, 40), WAM P.20069 (1, 45.5). Northern Territory: AMS IA.1556
(1, ca. 46.5) and AMS I.16288-001 (2, ca. 46-47.5), ‘“‘syntypes”’ of Ichthyocampus
annulatus. GCRL 16952 (1, 28), NIM S.10006-031 (1, 38), USNM 173071 (1,
44.5), USNM 173072 (1, 43.5), USNM 222931 (1, 30.5), USNM 222932 (1, 29.5).
Loc. uncertain: MNHN 84-897 (1, 45.5), Cochinchina.

Micrognathus natans, new species
Figs. 12, 13

Holotype.—AMS 1.20390-012 (36 mm SL, immature male), Fiji Is., Bega Is.,
probably taken at surface with dipnet and light, 14 Jan. 1974, B. Goldman and
B. Carlson.

Paratypes.—PHILIPPINE IS.: USNM 133055 (1, 33.5), S Luzon Is., Varadero
Hbr., 22 July 1908, Albatross. USNM 230674 (1, 33.5), S Luzon Is., Varadero
Bay, 20-23 July 1908, Albatross. USNM 230673 (1, 33.5), SE Mindoro Is., Man-
salay, 3 June 1908, Albatross. ZMUC P.39700 (1, 26), 13°32’N, 121°21’E, in 0-
100 m over 450 m, 26 June 1929, Dana Sta. 3733, HI. ZMUC P.39701 (1, 30) and
ZMUC P.39702 (1, 28), 11°43’N, 121°43’E, in 0-200 m over 1170 m, 27 June 1929,
Dana Sta. 3734, Il and II]. AUSTRALIA, Qld.: AMS 1.20951-007 (1, 26), Cape
York, E of Bligh Reef, 12°42’S, 144°05’E, neuston tow in 0-1 m over +900 m,
Sta. FNQ 79-82, 22 Feb. 1979, AMS-AIMS pty. AMS 1I.20909-001 (1, 36) and
GCRL 18237 (1, 45), Lizard Is. area, 16.7 km E of Carter Reef, 14°30’S, 145°52’E,
neuston tow in 0-1 m over +900 m, Sta. FNQ 79-14, 7 Feb. 1979, J. Leis and B.
Hartwick. AMS I.21760-001 (7, 34.5—45) and GCRL 18238 (5, 34.5—42.5), 11.1-
16.7 km E of Carter Reef, 14°30’S, 145°42’E, neuston tow in 0-1 m over +900
m, Sta. FNQ 79-113, 7 Feb. 1979, AMS-AIMS pty. AMS I.21761-001 (1, 37.5),
Sta. FNQ 79-114, data as for 79-113. AMS I.22540-001 (1, 37.5), Lizard Is.,
midway to outer barrier, 14°32'S, 145°34’E, 0-18 m, Sta. FNQ 79-7, 7 Feb. 1979,
AMS-AIMS pty. NEW CALEDONIA: AMS I.19762-039 (1, 34.5), ca. 71.4 km
offshore, 22°03’S, 167°44’E, midwater trawl in 0-800 m over 2300 m, 13 May
1971, J. Paxton. ZMUC P.39668-677 (10, 18.5—35.5), ZMUC P.39678-699 (22, 13-
30), GCRL 18136 (4, 20-31) and GCRL 18137 (7, 19-32), 20°53’12"S, 164°03’18"E,
in 0-50 m over 3490 m, 26 Nov. 1929, Dana Sta. 3611, V and VI.

VOLUME 95, NUMBER 4 683

Fig. 12. Micrognathus natans. Lateral and dorsal aspects of head and anterior trunk rings, to-
gether with section of body illustrating ridge configuration and dorsal and anal fins. From 42.5 mm
SL, female or juvenile male, paratype (GCRL 18238).

Diagnosis.—Trunk rings modally 14, principal ridges not angled laterad on
distal half of tail, dorsal-fin rays usually 17, subdorsal tail rings usually 3.75—5.0,
snout length averages 2.3 in HL, anal ring depth averages 4.8 in HL, side of
trunk without dark blotches above lateral ridge, opercle without dark spots or
irregular narrow bars.

Description.—Rings 14-15 + 27-31 (14 + 27-31 in 92%), dorsal-fin rays 16—
18, pectoral-fin rays 12-15 (modally 13), anal-fin rays 3—4 (usually 4), subdorsal
rings 0.5—0.0 + 3.5-5.0 = 3.75—5.0 (see Tables 1-3 for additional counts). Pro-
portional data, based on 10 specimens, 36.0-45.0 (« = 40.5) mm SL, follow: HL
in SL 6.9-7.8 (7.4), snout length in HL 2.2—2.5 (2.3), snout depth in snout length
2.9-4.0 (3.4), length of dorsal-fin base in HL 1.6—1.8 (1.7), pectoral-fin length in
HL 4.85.4 (5.1), length of pectoral-fin base in pectoral-fin length 1.4-2.0 (1.7),
anal ring depth in HL 4.3—5.2 (4.8), trunk depth in HL 3.1-3.7 (3.4). Measure-
ments (mm) of holotype (AMS I.20390-012) are: SL 36.0, HL 5.2, snout length
2.1, snout depth 0.7, length of dorsal-fin base 3.1, pectoral-fin length 1.0, length
of pectoral-fin base 0.6, anal ring depth 1.2, trunk depth 1.6. Longitudinal oper-

684 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 13. Top, Micrognathus natans, AMS 1.20390-012 (36 mm SL, immature male, holotype);
Bottom, M. erugatus. MZUSP 9408 (64 mm SL, presumed female, holotype).

cular ridge usually complete; subdorsal portion of superior trunk ridge usually
arched a little dorsad (Fig. 12), principal tail ridges not angled or flared laterad
(Fig. la), the posterior angles of rings not produced to hook-like points; dorsal-
fin origin often on tail, between anterior margin and middle of Ist tail ring in 24%
of specimens examined.

Holotype (Fig. 13) shaded or mottled with light-brown microchromatophores
on tan ground color; dorsum and upper half of side of trunk with irregular, diffuse,
pale bars on 4th, 8th and 12th rings; tail more or less encircled with similar
markings on 6th, 9th, 13th, 18th, 19th, 22nd and 26th rings. Distal margin and
basal third of dorsal fin hyaline; remainder of fin with a broad brownish stripe,
markings dark on anterior 8-10 rays but diffuse elsewhere. Pectoral and caudal
fins largely hyaline, anal-fin rays lined with brown. Coloration of other recently
preserved material similar to that of holotype, except for absence of the dorsal-
fin stripe and presence of brown shading on the caudal fin.

Comparisons.—Yhe combination of modally 14 trunk rings and 17 dorsal-fin
rays, long snout, and essentially straight tail ridges (Fig. la) distinguishes M.
natans from all congeners. It further differs in usually having fewer subdorsal
trunk rings (0.5—0.0 versus 1.75—0.25) and more pectoral-fin rays (12-15 versus
9-13) than its congeners. The snout length of M. natans most closely approaches
that of M. micronotopterus but the snout is more slender in M. natans (snout
depth in snout length averages 3.4 versus 2:6 in micronotopterus). Additionally,
M. natans has a lower average HL in SL ratio (7.4 versus 8.3) and somewhat
greater numbers of subdorsal tail rings (3.5-5.0 versus 2.25—3.75) than M. mi-
cronotopterus.

VOLUME 95, NUMBER 4 685

Remarks.—The holotype has a moderately long, slender, dermal flap near the
anterior end of the frontal ridge and there are short flaps located above the eye,
somewhat mesiad of the margin of each supraorbital ridge. The head bears other
single, short flaps on the opercular, prenuchal and nuchal ridges. The body has
short flaps, paired bilaterally, on the dorsum of the 2nd, 5th, 9th, 10th and 13th
trunk ring, as well as on the dorsum of some tail rings. A few minute flaps are
also present on the lateral trunk and inferior tail ridges. Among other material,
most of the larger specimens have some head or body flaps, but these are usually
absent from specimens smaller than about 35 mm SL.

The holotype is the only examined specimen with evidence of brood-pouch
development. This immature fish lacks pouch-protective plates but bilateral pouch
folds are developing on the venter of the anterior 10—11 tail rings.

Distribution.—Micrognathus natans is known from the Philippine Is., the
Queensland coast of Australia, the vicinity of New Caledonia and the Fiji Is.
(Fig. 5). Collection data for the holotype are incomplete but it was probably taken
at the surface with a nightlight and dipnet (B. Carlson, pers. comm.). Most of
the paratypes were taken with midwater trawl, neuston nets, or 2 m stramin nets
(Dana material) in the upper 0-800 m over depths of 450-3490 m.

Etymology.—Named from the Latin natans, swimming or floating, in view of
the occurrence of most, if not all, known specimens in the upper portions of the
water column.

Micrognathus erugatus Herald and Dawson
Fig. 13

Micrognathus erugatus Herald and Dawson, 1974:27, fig. | (orig. descr.; S of
Arembepe, Bahia, Brazil).

Diagnosis.—Trunk rings 20, principal tail ridges not angled laterad, snout length
3.1 in HL, anal ring depth 3.5 in HL.

Description.—Rings 20 + 36, dorsal-fin rays 19, pectoral-fin rays 13, anal-fin
rays 2, subdorsal rings 0.5 + 4.5 = 5.0. Measurements (mm) of holotype follow:
SL 64.0, HL 5.9, snout length 1.9, snout depth 1.0, length of dorsal-fin base 4.9,
pectoral-fin length 1.1, length of pectoral-fin base 0.7, anal ring depth 1.7, trunk
depth 1.9. Opercular ridge of holotype crosses about half of opercle length and
angles somewhat upward toward gill opening. Head with short, simple, dermal
flaps above opercle and on supraorbital and opercular ridges; trunk and tail with-
out dermal flaps.

Holotype (Fig. 13) dark brown, mottled with pale; trunk and tail with 15 diffuse
dark bars crossing dorsum and upper part of side; dorsal fin pale, except for
brown streaks near bases of fin-rays; caudal fin brownish with pale distal margin.

Comparisons.—This western Atlantic species is readily distinguished from Indo-
Pacific congeners by its high number of trunk rings (20 versus 14-17) and by
higher numbers of tail and total rings (Table 1). The essentially straight tail ridge
configuration of M. erugatus (Fig. la) is shared with three Indo-Pacific congeners
(andersonii, micronotopterus, natans) but it further differs from these species in
a number of proportional features (see descriptions).

Remarks.—Absence of spines on the head, absence of lateral snout ridge, pres-
ence of a low, entire, median dorsal snout ridge, together with close agreement
with congeners in most meristic and proportional features constitute reasons for

686 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

retention of this species in the genus Micrognathus. This decision is somewhat
tentative, since M. erugatus is presently known only from the holotype, and study
of additional material could result in reevaluation of the relationships of this
species. Occurrence of a western Atlantic species of a genus of pipefishes oth-
erwise restricted to the Indo-Pacific region is unusual but not without precedent.
The genus Penetopteryx Lunel is represented by P. nanus (Rosén) in the western
Atlantic and by P. taeniocephalus Lunel in the Indo-Pacific.

Distribution.—Known only from the Brazilian state of Bahia, southwestern
Atlantic Ocean.

Material examined.—Holotype, MZUSP 9408, 64 mm, presumed female, ca.
1 km S of Arembepe (ca. 12°45'S, 28°10'W), Bahia, Brazil, tidepool, 0-1.3 m, 26
Aug. 1972, C. E. Dawson and party.

Acknowledgments

I thank the curators and assistants of the referenced repositories for loans of
specimens and other courtesies. Special acknowledgment is due M. L. Bauchot
(MNHN), M. Boeseman (RMNH), W. N. Eschmeyer and Pearl Sonoda (CAS),
R. H. Gibbs, S. Weitzman and Susan Jewett (USNM), W. Klausewitz (SMF), J.
R. Paxton, D. F. Hoese and J. M. Leis (AMS) and A. C. Wheeler (BMNH) for
facilitating examination of type material in their care. Dr. P. J. Miller (University
of Bristol) provided several valuable specimens and detailed data from his Red
Sea collections; other gifts or exchange specimens were received from AMS,
CAS, USNM and the Smithsonian Oceanographic Sorting Center. Among GCRL
associates I thank Lois Coquet for computer related services, F. N. Jackson for
curatorial and photographic assistance, Yasue Matthews for drawings, and Eliz-
abeth Heal for contributions as Technical Secretary.

Literature Cited

Bleeker, P. 1853. Bijdrage tot de kennis der Troskieuwige visschen van den Indischen Archipel.—
Verhandelingen van het Bataviaasch genootschap van Kunsten en wetenschappen 25(6): 1-30.

—. 1858. Vijfde bijdrage tot de kennis der ichthyologische fauna van de Kokos-eilanden.—
Natuurkundig Tijdschrift voor Nederlandsch Indie 15:457-468.

Dawson, C. E. 1977. Synopsis of syngnathine pipefishes usually referred to the genus Len BEORE
pus Kaup, with description of new genera and species.—Bulletin of Marine Science 27(4):595—
650.

—. 1978. Review of the Indo-Pacific pipefish genus Bhanotia, with description of B. nuda n.
sp.—Proceedings of the Biological Society of Washington 91(2):392—407.

—. 1982. Descriptions of Cosmocampus retropinnis sp. n., Minyichthys sentus sp. n. and

Amphelikturus sp. (Pisces, Syngnathidae) from the eastern Atlantic region.—Zoologica Scripta

11:135-140.

, and G. R. Allen. 1981. Micrognathus spinirostris, a new Indo-Pacific pipefish (Syngnathi-

dae).—Journal of the Royal Society of Western Australia 64(2):65—-68.

, and R. A. Fritzsche. 1975. Odontoid processes in pipefish jaws.—Nature 257:390.

Dollfus, R. P., and G. Petit. 1938. Les Syngnathidae de la mer Rouge. Liste des especes avec la
description d’une sous-espece nouvelle.—Bulletin du Muséum de Histoire naturelle, Paris, 2°
serie 10(5):496—506.

Duncker, G. 1912. Die Gattungen der Syngnathidae.—Mitteilungen aus dem Naturhistorischen Mu-
seum in Hamburg 29:219-240.

—. 1915. Revision der Syngnathidae. Erster teil.—Mitteilungen aus dem Naturhistorischen
Museum in Hamburg 32:9-120.

VOLUME 95, NUMBER 4 687

Fishelson, L. 1971. Ecology and distribution of the benthic fauna in the shallow waters of the Red
Sea.—Marine Biology 10:113-133.

Fowler, H. W. 1938. Descriptions of new fishes obtained ae the United States Bureau of Fisheries
steamer “‘ALBATROSS,”’ chiefly in Philippine seas and adjacent waters.—Proceedings of the
United States National Museum 85:31-135.

Fritzsche, R. A. 1980. Revisions of the eastern Pacific Syngnathidae (Pisces: Syngnathiformes),

including both recent and fossil forms.—Proceedings of the California Academy of Sciences

42(6): 181-227.

1981. A new species of pipefish (Pisces: Syngnathidae: Micrognathus) from Tahiti.—Pro-
ceedings of the Biological Society of Washington 94(3):771-773.

Herald, E. S. 1953. Family Syngnathidae. In: L. P. Schultz et al., Fishes of the Marshall and:
Marianas Islands.—Bulletin of the United States National Museum 202(1):231-278.

——. 1959. From pipefish to seahorse—a study of phylogenetic relationships.—Proceedings of

the California Academy of Sciences, Fourth series 29(13):465—473.

, and C. E. Dawson. 1974. Micrognathus erugatus, a new marine pipefish from Brazil

(Pisces: Syngnathidae).—Proceedings of the Biological Society of Washington 87(4):27—30.

, and J. E. Randall. 1972. Five new Indo-Pacific pipefishes.—Proceedings of the California
Academy of Sciences, Fourth series 39(11):121—140.

Jenyns, L. 1842. Fish. In The Zoology of the Voyage of H.M.S. BEAGLE, under the command of
Captain Fitzroy, R. N., during the years 1832 to 1836. Smith, Elder and Co., London, 4:1-172.

Jordan, D. S., and E. C. Starks. 1906. List of fishes collected on Tanega and Yaku, offshore islands
of southern Japan, by Robert Van Vleck Anderson, with descriptions of seven new species.—
Proceedings of the United States National Museum 30:695-706.

Kaup, J. 1853. Uebersicht der Lophobranchier.—Archiv fur Naturgeschichte, Berlin 19:226—234.

1856. Catalogue of lophobranchiate fish in the collection of the British Museum.—Taylor
and Francis, London, 76 pp.

-Klunzinger, C. B. 1871. Synopsis der Fische des Rothen Meeres.—Verhandlungen der Kaiserlich-
koniglichen zoologisch-botanischen Gesellschaft in Wien 21:441-688.

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Wales 2:344-367.

Marshall, N. B. 1950. Fishes from the Cocos Keeling Islands.—Bulletin of the Raffles Museum 22:
166-205.

Ogilby, J. D. 1908. New or little known fishes in the Queensland Museum.—Annals of the Queens-
land Museum 9(1):1—41.

Ruppell, E. 1838. Neue Wirbelthiere zu der Fauna von Abyssinien geéhorig.—Frankfurt am Main,
Lieferung 12:81-148.

Smith, J. L. B. 1955. The fishes of Aldabra.—Part I—Annals and Magazine of Natural History,
12th series 8:304—312.

——. 1963. Fishes of the family Syngnathidae from the Red Sea and the western Indian Ocean.—

Rhodes University Ichthyological Bulletin 27:515—543.

, and M. M. Smith. 1963. The fishes of Seychelles.—Department of Ichthyology, Rhodes

University, Grahamstown, 215 pp.

Springer, V. G., and W. N. Eschmeyer. 1974. Location of holotypes of Japanese fishes (Jordan
and Starks, 1906) and types of Hawaiian fishes (Jordan and Evermann, 1903).—Copeia 1974(2):

566-568.

Weber, M. 1913. Die Fische der Siboga-Expedition.—Siboga Expedition monograph 57:i-xii + 1-
710.

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84 pp.

Gulf Coast Research Laboratory Museum, Ocean Springs, Mississippi 39564.

PROC. BIOL. SOC. WASH.
95(4), 1982, pp. 688-693

TAXONOMY OF BUFO VENUSTUS PHILIPPI, 1899
(ANURA: LEPTODACTYLIDAE) FROM CENTRAL CHILE

J. R. Formas and Alberto Veloso

Abstract.—The frog Bufo venustus Philippi, 1899, is shown to be a species of
the genus Telmatobufo but not conspecific with T. bullocki. The species is re-
described, and some notes on its natural history are presented.

Philippi (1899) described Bufo venustus on the basis of three frogs collected in
Cordillera de Chillan (Andes Range, Chillan Province), and one frog from San
Ignacio de Pemehue (Cautin Province), Chile. Donoso-Barros (1972) without os-
teological analysis of the holotype suggested that this species should be trans-
ferred to the genus Aruncus Philippi, 1902, and considered Telmatobufo bullocki
Schmidt, 1952, conspecific with A. venustus. Lynch (1978) placed the species in
the genus Telmatobufo, but the frogs (KU 159811 and 161439) analyzed by Lynch
were true T. bullocki rather than venustus.

Recently one frog with external similarity (habitus and color pattern) to Philip-
pi’s description of Bufo venustus was collected in Alto Vilches, Chile (Talca
Province, 1280 m). The color of the holotype could not be checked by Donoso-
Barros (1972) because its original color is lost (it is whitish in alcohol).

In order to establish the taxonomic status of B. venustus we analyzed the
osteology (skull and pectoral girdle) of the holotype (MUZUC 205051) and the
Alto de Vilches’ specimen (DBG 0784) as well as the external morphology of
both frogs. Characteristics were compared with those of Telmatobufo bullocki
and with Telmatobufo australis Formas, 1972.

As result of the study we concluded that Bufo venustus belongs to the genus
Telmatobufo because the paratoid glands are well developed, the pectoral girdle
is arciferal with the omosternum short and the sternum broad. The frontoparietals
are nearly meeting at the midline, the nasal bones are medium-sized and broadly
separated. The maxillary arch is complete and the cultriform process of the para-
sphenoid reaches the prevomerine teeth. Telmatobufo bullocki Schmidt, 1952, is
not conspecific with Philippi’s species, differing in color pattern and morphology
of the foot.

Telmatobufo venustus (Philippi, 1899)
Fig. |

Bufo venustus.—Philippi, 1899:723. Chile: Cordillera de Chillan.
Aruncus venustus.—Donoso-Barros, 1972: 109-116.
Telmatobufo venustus.—Lynch, 1978:24—27.

Holotype.—MUZUC 205051, an adult female from Cordillera de Chillan, Chil-
lan Province, Chile, approximately 1200 m, collected in January 1897 by Germain.

Diagnosis.—Telmatobufo venustus is a moderate size frog that is distinguished
from its congeners in having dorsal and ventral areas black, the dorsum, head,

VOLUME 95, NUMBER 4 689

Fig. 1. Living specimen (above) and holotype (below) of Telmatobufo venustus.

and paratoid glands spotted with orange, and the outer border of the fifth toes
broad and not glandular.

Description.—This redescription is based on two specimens, the holotype and
one juvenile (DBG 0784). Body robust, limbs long and slender. Head depressed,
slightly wider than long; head length 33.2% of snout—vent length; snout short,
truncate (slightly concave in holotype). Lateral nostrils protruding; nostrils closer
to the tip of snout than to eye. Eye length greater than distance between eye and
nostril; internarial distance greater than interorbital distance. Tympanum absent;
paratoid glands large, oval, greater than diameter of eye. Tongue rounded; pre-
vomerine teeth in two separated fascicules between large choanae.

690 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

b

Fig. 2. Foot of (a) T. bullocki and (b) T. venustus. Scale = 5 mm.

Posterior limbs thin; tibiotarsal articulation reaching to anterior edge of paratoid
gland (in DBG 0784). Toes webbed (Fig. 2b), interdigital membrane extending to
tips of toes; length of toes in decreasing order 4, (S—3) 2, 1; inner metatarsal
tubercle flat, indistinct; outer metatarsal tubercle absent; outer border of the fifth
toe broad and not glandular; subarticular tubercles indistinct; tarsal fold present.

Forelimbs thin; upper arm enveloped in folds of skin of body; fingers free, their
lengths in decreasing order 3—4—2—1; inner palmar tubercle evident; elongated
gland in ulnar position of forearm extending from elbow to hand.

Skin of dorsum with numerous prominent, oval and round glands (less prom-
inent in holotype); skin of venter smooth.

Color in preservative, whitish in holotype and black with irregular whitish spots
in the juvenile. In life this animal had the venter, dorsum and extremities bright
coal-black; head, paratoid glands, upper eyelids, elbow and tibiotarsal articula-
tion areas orange spotted; two orange paravertebral stripes convergent below
paratoid glands; minute yellow spots on flanks, dorsal and posterior thigh, dorsal
surface of tarsus, and inner edge of forearm; tips of fingers and toes yellow. The
color pattern of the living DBG 0784 here described agrees with Philippi’s de-
scription of the holotype. °

Osteology.—Pectoral girdle arciferal. Clavicles arched and not in contact me-

VOLUME 95, NUMBER 4 691

Table 1—Measurements (mm) of Telmatobufo venustus, T. bullocki, and T. australis.

T. venustus T. australis
MUZUC 205051 T. bullocki IZUA 934-A
DBG 0784 n= 11 IZUA 1778-A
Holotype Juvenile aye Range x + SD Holotype Juvenile
Snout—vent length 70.8 46.9 80.0-58.4 697] s== el 40.0 70.6
Head width 24.9 19.4 29.5—23.8 Sod) se DLA 18.0 27.0
Head length W335) 17.6 28 12a Dest DAO) 19.0 23.0
Thigh length 28. 1 DRA 38.0-25.4 82) ae Gl.5 18.0 33.4
Shank length 29.4 OND 37.0-28.0 31.0 + 3.8 16.0 30.8
Foot length 46.5 31.9 61.1-46.8 52.4 + 9.0 21.0 SoZ
Paratoid length 10.6 7.6 14.0-8.5 10.2 + 1.6 6.1 11.0
Eye diameter 8.3 6.2 9.8-7.1 8.5 + 0.9 58) 8.2
Eye-—nostnil distance 28) 3)a8) 5.34.6 49+ 0.2 37) 53)
Nostril-snout distance 5.3 wih 11.2—3.5 6.4 + 3.0 3.9 Soo)
Internarial width Vel Sal 7.0-6.3 6.7 + 0.3 4.9 8.3
Interorbital width 6.5 4.4 8.0-6.5 6.8 + 0.6 4.4 7.0

dially, coracoids expanded at distal and proximal ends; epicoracoidal cartilages
free and broad; sternum very broad, slightly notched; omosternum broad and
short, not ossified as sternum.
_ Cranial osteology.—The skull of the holotype of T. venustus and DBG 0784
were studied by radiographs. Frontoparietals paired, nearly meeting along the
midline, sutured posteriorly and resting on the otoccipital; anteriorly resting on
the sphenethmoid. Nasal bones medium-sized and broadly separated, the max-
illary processes not contacting the maxillae; margins of nasals resting on antero-
lateral margin of sphenethmoid. Sphenethmoid extending slightly anterior to na-
sals. Maxillary arch complete and quadratojugal of moderate size. Zygomatic
ramus of squamosal long, otic ramus shorter. Parasphenoid triradiate, cultriform
process long, reaching prevomerine teeth. Pterygoids small and strong, their me-
dian rami short. Palatines broad, extending onto edge of sphenethmoid. Prevo-
mers medium-sized, broadly separated, bearing 3-4 fang-like teeth. The skull of
this species has the same osteological characteristics described by Lynch (1978)
for T. venustus (=bullocki).

Comparisons

Telmatobufo venustus shows a remarkable color pattern which is different from
T. bullocki and T. australis. In preservative (alcohol) the dorsum is brown in T.
bullocki, and gray in T. australis. The venter is clear gray in 7. australis with
dark irregular spots on a whitish background (Formas 1972, 1979); T. bullocki
shows this area mottled with brown (Schmidt 1952). In life, orange spots and
stripes are absent in 7. australis and T. bullocki. The latter species always has a
yellow stripe between the eyes and some specimens show a yellow reticulate
pattern among the warts (Pefaur 1971). In life the juveniles of 7. australis are
dark green with two yellow paravertebral stripes (Formas 1972).

The outer side of the fifth toe of 7. venustus is broader than in T. bullocki and
no glandular areas are present. Telmatobufo australis does not show this glan-

692 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

dular fold on the outer side of this toe. The tips of the toes of T. bullocki and T.
australis are pointed, whereas in 7. venustus they are rounded (Fig. 2a, b). The
cranial osteology of 7. venustus agrees with the description of the skull given by
Lynch (1978) for T. bullocki (hU 159811), and the pectoral girdle agrees with
Schmidt’s (1952) illustration. Formas found the same cranial patterns and pectoral
girdle morphology in 7. australis.

Measurements of 7. venustus, T. bullocki, and T. australis are given in Table 1.

Natural History and Distribution

Only four specimens of 7. venustus were obtained by Philippi, one from San
Ignacio de Pemehue (Cautin Province) and three from the Cordillera de Chillan.
The newly collected specimen from Alto de Vilches (the northernmost locality
from which 7. venustus is known) was found under a Nothofagus log on 10
September 1979. The following food items were identified in DBG 0784: one snail,
one adult coleopteran, one adult and one larval of dipteran, one adult hemipteran,
and plants. One of us (Veloso) observed that its swimming activity consisted of
asynchronic movements of the extremities, like those described by Donoso-Bar-
ros (1972) for T. bullocki. In the laboratory, the frog remained with the nares
outside the water and never was observed immersed. The eggs are creamy-white
in color. The paucity of collected specimens of this species suggests to us that
the populations of this frog are small.

Specimens Examined

Abbreviations.—Instituto de Zoologia, Universidad Austral (IZUA); Museo de
Zoologia Universidad de Concepcion (MUZUC); University of Kansas, Museum
of Natural History (KU); Chicago Field Museum of Natural History (CNHM);
Departamento de Biologia Celular y Genética, Universidad de Chile (DBG).

Telmatobufo bullocki: CHILE: CNHM 23842, 31618, Cerros de Nahuelbuta,
Malleco Province; KU 18438, 159811, Los Lleulles, Malleco Province. MUZUC
11644, Lota, Concepcion Province; 12276, Elicura, Concepcion Province; 12266,
Ramadillas, Arauco Province; IZUA 1563, Ramadillas, Arauco Province; 1858-
9, Caramavida River, Arauco Province. DBG, uncatalogued, locality imprecise.

Telmatobufo venustus: CHILE: MUZUC 205051, Cordillera de Chillan, Chillan
Province; DBG 0784, Alto de Vilches, Talca Province, 1280 m.

Telmatobufo australis: CHILE: IZUA 934-A, 1778-A, Cordillera Pelada, Val-
divia Province.

Acknowledgments

We are grateful to Francisco Silva and Carlos Vivar (Universidad de Valpa-
raiso) who collected the specimen (DBG 0784), and Jorge Artigas (Universidad
de Concepcion) for making specimens available for our study. The first author
wishes to express special thanks to William E. Duellman who provided facilities
and made specimens available during his stay at the University of Kansas. Sonia
Lacrampe and M. Inés Vera gave us technical assistance. This study was partially
supported by the Direccion de Investigacion y Desarrollo, Universidad Austral
de Chile (Proyecto S-79-3).

VOLUME 95, NUMBER 4 693

Literature Cited

Donoso-Barros, R. 1972. Contribucién al conocimiento del género Aruncus Philippi.—Boletin So-
ciedad Biologia Concepcion 44:109-116.

Formas, J. R. 1972. A second species of Chilean frog genus Telmatobufo (Anura: Leptodactyli-
dae).—Journal Herpetology 6:1-3.

——. 1979. New observations of Telmatobufo australis (Anura, Leptodactylidae) in Southern
Chile.—Journal Herpetology 13:359-361.

Lynch, J. 1978. A re-assessment of the telmatobiine leptodactylid frogs of Patagonia.—Occasional
Paper, Museum Natural History, University of Kansas 72:1-—57.

Pefaur, J. 1971. Nota sobre Telmatobufo bullocki (Anura, Leptodactylidae).—Boletin Museo His-
toria Natural Chile 32:215—225.

Philippi, R. A. 1899. Descripciones breves de dos nuevas especies de sapos (Bufo).—Anales Uni-
versidad de Chile 104:723-725.

—. 1902. Suplemento a los Batraquios Chilenos descritos en la Historia Fisica y Politica de
Chile de don Claudio Gay.—Santiago, pp. 1-161.

Schmidt, K. P. 1952. A new leptodactylid frog from Chile.—Fieldiana Zoology 34:11-15.

(JRF) Instituto de Zoologia, Universidad Austral de Chile, Casilla 567, Valdi-
via, Chile. (AV) Departamento de Biologia Celular y Genética, Universidad de
Chile, Casilla 6556, Santiago, Chile.

PROC. BIOL. SOC. WASH.
95(4), 1982, pp. 694-701

GYMNODORVILLEA FLORIDANA, A NEW GENUS AND
SPECIES OF DORVILLEIDAE (POLYCHAETA)
FROM SOUTHEASTERN FLORIDA

Sam C. Wainright and Thomas H. Perkins

Abstract.—Gymnodorvillea floridana, n. gen., n. sp. (Polychaeta: Dorvillei-
dae), is proposed for specimens collected off southeastern Florida. Gymnodor-
villea appears to be a continuation of the Schistomeringos-Pettiboneia line of
dorvilleid evolution. The principal diagnostic characters are the complete lack of
prostomial appendages and dorsal cirri, and a unique jaw apparatus.

During an investigation of polychaete-sediment relationships in shallow waters
off Highland Beach, Palm Beach County, Florida (Wainright, in prep.), it was
found that the most common species of Dorvilleidae is undescribed and possesses
characters not assignable to any currently accepted genus.

Benthic core samples were collected in summer and winter during 1978 at 5
stations along a transect in depths of 3.7-19.8 m. Stations 4 and 5 yielded spec-
imens of the new species.

Station 4 (26°24.6’N, 80°03.1'W) was located near a 3—5 m diameter patch reef
approximately 20-30 m shoreward of the major reef tract parallelling the shoreline
of Dade, Broward, and Palm Beach Counties, Florida. Mean depth was 19.8 m;
sediments were soft, white to grey, moderately-sorted, fine-grained, calcar-
eous-siliceous sand. Collections were made on 28 February and 27 August 1978.
Station 5 was located at the same coordinates and depth as Station 4, but adjacent
to the major reef tract noted above. Sediments were moderately- to poorly-sorted,
medium- to fine-grained. Collections were made on 23 February and 20 August 1978.

Samples were fixed immediately after collection in 15% seawater formalin with
Rose Bengal dye added, and sieved about 24 hrs later using a 1 mm screen. The
remaining material was transferred to 70% ethanol.

Six specimens were serially transferred to 100% ethanol, critical-point dried in
CO., coated with 3 x 125 A gold-palladium and photographed with an ISI Super
III A scanning electron microscope. Four specimens were mounted on slides
using CMC-10 clearing-mounting medium for examination of mouthparts. Exter-
nal body parts were drawn with aid of a camera lucida attachment on a Wild M20
microscope or from SEM photomicrographs; mouthparts were drawn using a
camera lucida and Zeiss Nomarski interference-contrast optics.

Types and additional specimens are deposited in the National Museum of Nat-
ural History, Smithsonian Institution (USNM), the Florida Department of Natural
Resources, Marine Research Laboratory (FSBC I), and the Department of Bio-
logical Sciences, Florida Atlantic University (FAU).

Anatomical nomenclature follows that of Perkins (1979). Nomenclature of
mouthparts follows Jumars (1974). Free denticles are numbered sequentially from
the posterior end as in Perkins (1979).

VOLUME 95, NUMBER 4 695

Gymnodorvillea, new genus

Type-species.—Gymnodorvillea floridana, new species.

Diagnosis.—Body minute, slender, cylindrical, with about 20-30 setigers; pro-
stomium slightly flattened, pear-shaped, lacking antennae and palps, having 2
encircling bands of cilia; parapodia uniramous, without notoacicula and dorsal
cirri, with ventral cirri beginning on setiger 2; setae including serrate capillary,
furcate, geniculate, and cultriform simple setae and heterogomph falcigers; jaws
consisting of mandibles and 2 rows of maxillary denticles, with reduced basal
plates; carriers reduced if present; denticle rows fused posteriorly; denticles of 3
types: rounded serrate, serrate with principal tooth, and spinous; mandibles with-
out free lateral teeth.

Etymology.—The generic name is derived from Greek, gymno meaning naked
plus Dorvillea and refers to the lack of prostomial appendages. Gender: feminine.

Gymnodorvillea floridana, new species
Figs. 1-4

Material examined.—Highland Beach, Wainright et al., collectors; Sta. 4, win-
ter, holotype (USNM 71445), 9 paratypes (USNM 71446; FSBC I 28986); Sta. 4,
summer, 3 paratypes (USNM 71447; FSBC I 28987; FAU); Sta. 5, winter, 9
paratypes (USNM 71448; FSBC I 28988; FAU); Sta. 5, summer, 5 paratypes
-(USNM 71449; FAU).

Description.—Body (Fig. 1A) without color pattern in alcohol; largest specimen
2.9 mm long, 0.24 mm wide. Prostomium without eyes, without appendages. Two
apodous peristomial rings, each with transverse ciliated band at least dorsally
and laterally; length of anterior ring about '2—%4 that of posterior ring. Dorsal,
transverse ciliary bands from first peristomial ring to about setiger 14 (Fig. 2).

Parapodia supported by | or 2 neuroacicula (Fig. 1B); upper one resembling
single aciculum of other parapodia, stout, thicker than setal shafts, tapering,
sometimes protruding through parapodial lobe; lower aciculum apparently absent
from some parapodia, more slender, not protruding, located below cluster of setal
shafts surrounding upper aciculum. Presetal and postsetal lobes similar, indistinct
(Fig. 1B—E). Acicular lobes often protruding; lower acicular lobe protruding es-
pecially in parapodia with cultriform setae (Fig. 1B, E). Ventral cirri bluntly
conical, arising about midway along parapodium (Fig. 1D), extending ventrolat-
erally about 4—'4 parapodial length (Fig. 1C—E).

Setae of 5 types: (1) furcate setae with about 3 rows of serrations on short
prongs (Figs. II, 2); (2) geniculate setae (Fig. 1H); (3) serrated capillaries (Figs.
IF, G); (4) heterogomph falcigers with blades having serrate cutting edges and
entire tips, with blade lengths decreasing dorsally to ventrally, with blades joined
with shafts by thin membranes, with shafts having 2—3 rows of serrations below
joints (Figs. 1J—L); (5S) simple serrate cultriform setae (Fig. 1M).

Furcate setae supra-acicular, beginning on setiger | (28 specimens) or setiger
2 (4 specimens), rarely 2 per parapodium. Geniculate setae replacing furcate setae
in middle or posterior segments (range for setiger of replacement: 2-29, Y = 12.8,
SD = 8.0, n = 19), occasional furcate seta occurring posterior to first appearance
of geniculate setae, geniculate and furcate setae rarely occurring on same para-

696 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Gymnodorvillea floridana, paratypes: A, Dorsal view, outer margins of ciliary bands on
prostomium and apodous rings shown, bands on subsequent segments omitted; B, 11th parapodium,
ventroposterior view, showing acicula; C, 14th parapodium, posteroventral view; D, 11th parapodium,
posterior view; E, 20th parapodium, anterior view; F, Capillary seta; G, Portion of same, edge-on
view; H, Geniculate seta; I, Furcate seta; J, Upper falciger; K, Middle falciger; L, Lowest falciger;
M, Cultriform seta. (CE, G, I, J drawn from SEM photomicrographs.)

podium. Serrate capillaries appearing supra-acicular on parapodia with furcate
setae, subacicular on parapodia with geniculate setae, 1, occasionally 2, per para-
podium. Subacicular falcigers occurring in fascicles of 3 on anterior parapodia, 2
on middle or posterior parapodia, but often 4 or 3, respectively. Cultriform setae
beginning on setiger I-11 (Y = 6.3, SD = 3.6, n = 19) similar to and replacing
lower falcigers.

Mandibles (Fig. 3A) fused, flared, and thinly chitinized anteriorly, each with
small medial tooth just anterior to point of fusion, posteriorly curved, forming
characteristic horseshoe shape and more thickly chitinized. Maxillae (Figs. 3B,
4A, B) forming single pair of denticle rows. Carriers, if present, greatly reduced,
formed of thin, dorsolateral, winglike structures fused to basal parts of maxillary

VOLUME 95, NUMBER 4 697

Fig. 2. Gymnodorvillea floridana, paratypes: A, Anterior region, dorsal view, showing ciliary
bands, shrinkage due to fixation and/or critical-point drying; B, Furcate seta (SEM photomicrographs.
Scales: A = 50 um, B = 5 um).

rows (Figs. 3Bb, 4A, B), with points of fusion indistinct; parts of ligaments of D
I-5 attached to winglike structures on basal pieces; posterior pieces of denticle
rows fused, continuous with posterior tonguelike projection of ligament (Fig.
3Ba); anterior central part of ligament darkened. Remnants of basal plates prob-
ably consisting of posterior maxillary piece and D 1-5 (Fig. 4B). Denticles of 3
distinct types: (1) D 1| (Fig. 3Bc) ventrally hollow, with row of about 9-15 irregular
serrations; (2) D 2-5 ventrally hollow, with single, falcate major tooth directed
dorsoposteriorly with row of fine serrations on border, with serrations on medial
sides beginning with D 3 and increasing anteriorly; D 5 with anterior margin
elongate dorsoanteriorly (Fig. 4A); (3) D 6-8, spinous, padlike, with rounded
Surface covered with numerous fine spines (spines much more numerous than
indicated on figures); D 6 approximately same size as D 5, with reniform spinous
surface directed medially when pharynx withdrawn (Fig. 4); D 7 oval, with spi-
nous pad, extending anteriorly as indistinct chitinous plate; D 8 overlapping D 7
laterally and dorsally, with hemispherical spinous surface, beaklike posterior pro-
jection, and indistinct anterior plate (Fig. 3B).

698 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

an
TATA

Ein,
4 (4
GEG
LG

7

2 typ,
Wavy

0.025mm

Fig. 3. Gymnodorvillea floridana, paratypes: A, Mandibles, dorsal view; B, Maxillae, denticles
1-6 turned, posterior maxillary piece broken, a, posterior tonguelike extension of maxillary ligament,
b, winglike projection on basal piece, c, posterior denticle.

Pygidium truncate, with 2 clubshaped anal cirri.

Sexually mature specimens with gametes in about medial 10 setigers. Eggs of
one specimen measuring 3—5 jm in diameter.

Etymology.—The specific name refers to the type locality.

Biology.—Specimens were collected only at the 19.8 m stations, although three
Sshallower-water stations (3.7, 7.6, and 12.2 m) were sampled, suggesting a re-
striction to deeper water or to areas near coral reefs. The gut of one specimen,
viewed through the body wall, contained what appeared to be diatom chains,
agreeing with the feeding mode described by Fauchald and Jumars (1979) for the

VOLUME 95, NUMBER 4 699

Fig. 4. Gymnodorvillea floridana, paratypes: A, Maxillae, dorsal view, turned; B, Same, of smaller
paratype, only very slightly turned.

family. The greater number of specimens collected in winter samples at both
Stations suggests a fall or winter reproductive periodicity. Depth of occurrence
within the sediment could not be determined, although lack of head appendages
and eyes and minute body size imply an infaunal existence.

Remarks.—Currently accepted dorvilleid genera were defined by Jumars (1974),
Blake (1979), and Gaston and Benner (1981); however, characters of species
described in recent papers by Oug (1978) and Armstrong and Jumars (1978) in-
dicate that a generic revision of the family is needed.

Gymnodorvillea appears most closely related to Pettiboneia Orensanz (1973),
both genera being allied with Jumars’ (1974) Schistomeringos-Protodorvillea-
Meiodorvillea line of dorvilleid evolution. Both genera have similar maxillary

700 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

denticles. Maxillae of Pettiboneia are organized into 7 pairs of denticle rows,
whereas Gymnodorvillea has only a single pair. The denticles of the dorsalmost
row of Pettiboneia appear identical to the falcate, proximal denticles of Gym-
nodorvillea, and denticles of the remaining rows of Pettiboneia appear similar to
the distal three pairs of spinous denticles of Gymnodorvillea. Thus it appears that
maxillae of Gymnodorvillea have evolved into a single pair of denticle rows through
fusion of the dorsal rows of falcate denticles with one of the paired rows of
spinous maxillary denticles of a Pettiboneia-like ancestor. Also, except for the
absence of geniculate setae, the setae of Pettiboneia are similar to those of Gym-
nodorvillea. Conversely, Gymnodorvillea differs from Pettiboneia in lacking palps,
antennae, notoacicula, and dorsal cirri, and in the fusion of posterior denticles,
which are well separated in Pettiboneia. However, notoacicula and dorsal cirri
are slightly reduced on anterior segments and absent on posterior segments of
Pettiboneia, and antennae and palps are reduced, further indicating that Gym-
nodorvillea is closer to Pettiboneia than to other genera (except perhaps Meio-
dorvillea Jumars). Additionally, some parapodia of Gymnodorvillea have a sec-
ond neuroaciculum ventral to the principal one. The stouter, upper aciculum is
apparently homologous with the solitary one of most other genera. The ventral
one may be a development analogous to the ventral aciculum found in the dor-
villeids Ophryotrocha lobifera Oug, 1978, and Meiodorvillea apalpata Jumars,
1974.

It is also suggested that Protodorvillea gaspeensis Pettibone (1961:178, 179)
should be placed near the Schistomeringos-Pettiboneia-Gymnodorvillea line of
dorvilleid evolution. The species lacks the distinct maxillary carriers found in
Protodorvillea Pettibone; the fused, posterior maxillary pieces of P. gaspeensis,
considered to be carriers by Jumars (1974) and thus a remnant of a third pair of
maxillae, are more simply characterized as fused posterior pieces of the dorsal
denticle rows. Mandibles and maxillary denticles of P. gaspeensis are similar to
those of Pettiboneia and Gymnodorvillea, and are especially similar to those of
Pettiboneia in that they have repeating rows of spinous denticles (based on recent
observations by T.H.P. of specimen figured by Jumars 1974:118, Fig. 8; USNM
43516). However, P. gaspeensis cannot be assigned to Pettiboneia because it
lacks notopodial cirri and acicula, and because the posterior maxillary pieces of
the upper denticle rows are joined rather than well separated. Further, since both
prostomial appendages and repeating rows of denticles are present, P. gaspeensis
should probably not be assigned to Gymnodorvillea, but instead to a new genus.

Gymnodorvillea is similar to Meiodorvillea Jumars, 1974, in having maxillae
with only 2 rows of denticles, uniramous parapodia, and furcate or geniculate
setae, but the latter differs in having distinct maxillary carriers, antennae, and
usually palps. Meiodorvillea apalpata Jumars, which lacks palps, also has para-
podia with a second neuroaciculum and may prove to belong to another genus.

The position and distribution of furcate and geniculate setae on the parapodia
of Gymnodorvillea floridana suggests that they are homologous. Also, in our
opinion, the posterior denticles (D 1) which do not have a large principal tooth
are homologous with those denticles which have such a tooth (D 2-5). These are
not considered to be important generic characters.

An alternative interpretation to that presented in the description for the max-
illary organization of Gymnodorvillea floridana may be that the proximal denticles

VOLUME 95, NUMBER 4 701

(D 1) and the fused posterior maxillary pieces together constitute the basal plates,
as suggested by Fig. 4. |

Acknowledgments

Dr. George T. Taylor graciously furnished the use of Florida International
University’s scanning electron microscope and spent a generous amount of time
instructing the senior author in its use. Dr. G. Alex Marsh constructively criti-
cized the manuscript. Material costs were partially paid by Florida Atlantic Uni-
versity.

Literature Cited

Armstrong, J. W., and P. A. Jumars. 1978. Branchiate Dorvilleidae (Polychaeta) from the North
Pacific.—Bulletin of the Southern California Academy of Sciences 77(3): 133-138.

Blake, J. A. 1979. A redescription of Pettiboneia sanmatiensis Orensanz (Polychaeta: Dorvilleidae)
and a revised key to the genera of the Dorvilleidae.—Bulletin of the Southern California Acad-
emy of Sciences 78(2): 136-140.

Fauchald, K., and P. A. Jumars. 1979. The diet of worms: A study of polychaete feeding guilds.—
Oceanography and Marine Biology; An Annual Review 17:193—284.

Gaston, G. R., and D. A. Benner. 1981. On Dorvilleidae and Iphitimidae (Annelida: Polychaeta)
with a redescription of Eteonopsis geryonicola and a new host record.—Proceedings of the
Biological Society of Washington 94(1):76-87.

Jumars, P. A. 1974. A generic revision of the Dorvilleidae (Polychaeta), with six new species from
the deep North Pacific.—Zoological Journal of the Linnean Society of London 54:101—135.

Orensanz, J. M. 1973. Los anélidos poliquetos de la provincia biogeografica Argentina III. Dorvil-
leidae.—Physis, Buenos Aires, Seccion A. 32(85):325—342.

Oug, E. 1978. New and lesser known Dorvilleidae (Annelida, Polychaeta) from Scandinavian and
northeast American waters.—Sarsia 63(4):285—303.

Pettibone, M. A. 1961. New species of polychaete worms from the Atlantic Ocean, with a revision
of the Dorvilleidae.—Proceedings of the Biological Society of Washington 74: 167-186.
Perkins, T. H. 1979. Lumbrineridae, Arabellidae, and Dorvilleidae (Polychaeta), principally from
Florida, with descriptions of six new species.—Proceedings of the Biological Society of Wash-

ington 92(3):415—466.

Wainright, S. C. (In preparation). M.S. Thesis, Florida Atlantic University, Boca Raton, Florida.

(SCW) Department of Biological Sciences, Florida Atlantic University, Boca
Raton, Florida 33431; (THP) Florida Department of Natural Resources Marine
Research Laboratory, 100 Eighth Avenue SE, St. Petersburg, Florida 33701.

[Direct reprint requests to THP.]

PROC. BIOL. SOC. WASH.
95(4), 1982, pp. 702-708

DESCRIPTION OF DACTYLOKEPON SULCIPES N. SP.
(CRUSTACEA: ISOPODA: BOPYRIDAE) AND
NOTES ON D. CARIBAEUS

Daniel L. Adkison

Abstract.—Dactylokepon sulcipes, n. sp. is described from material collected
in the eastern Gulf of Mexico on the leucosiid crab, Callidactylus asper Stimpson.
Dactylokepon sulcipes is most similar to D. caribaeus. The female of D. sulcipes
differs from that of D. caribaeus by 1) having different ornamentation of the
internal ridge of the first oostegite; 2) having a posterolateral point on the first
oostegite; 3) having a groove on the external (ventral) surface of the fifth ooste-
gite, and 4) having shorter endopods on the pleopods. The male of D. sulcipes
has midventral tubercles on the pereon and pleon; the male of D. caribaeus lacks
these tubercles.

Among the crabs collected by Dr. Thomas S. Hopkins from the eastern Gulf
of Mexico were four Callidactylus asper Stimpson, 1871, having prominent bran-
chial swellings indicating infestation by a bopyrid isopod. These crabs and their
bopyrids were given to the author for study. The bopyrids were first thought to
be Dactylokepon caribaeus Markham, 1975. Upon examination of the types at
the Smithsonian Institution, it became apparent that the bopyrid from C. asper
represented an undescribed species.

The description of D. caribaeus is supplemented with a few observations that
aid in distinguishing these two bopyrids.

Dactylokepon caribaeus Markham, 1975
Fig. 1

Daktylokepon (sic) caribaeus Markham, 1975:61.
Dactylokepon caribaeus Markham, 1975:61-—66, figs. 4—6.

Material examined.—Infesting Iliacantha subglobosa Stimpson, 1871. Off-
shore southeast coast of Dominican Republic; Pillsbury station P-1387; 18°21'N,
69°09'W;, 130-165 m; 9 July 1971; 1 2 (paratype) USNM 143657, | ¢ (allotype)
USNM 143656. Same general locality; Pillsbury station P-1395; 18°21’N, 69°12'W;
166 m; 10 July 1971; | 2 (holotype) USNM 143654, | ¢ (paratype) USNM 143655.

Description.—Only points differing from or not mentioned by Markham (1975)
are given.

Female: (Fig. 1). Few tubercles present on pereomeres 1-4, particularly tergal
area. Dorsal bosses with lateral margins crenulate. Coxal plates present on all
pereomeres, greatly reduced on posterior pereomeres. Oostegite 5 without ridge
and groove proximally. Pleopods with endopods nearly length of exopod, ratio
of endopod to exopod not decreasing posteriorly.

Male: as described by Markham (1975).

Distribution.—Known from the Caribbean Sea off the southeastern coast of

VOLUME 95, NUMBER 4 703

Q)
= “4
A)
<_< C~
\

Fig. 1. Dactylokepon caribaeus, female, holotype: a, Left lateral plate and pleopod 4; b, Left
lateral plate and pleopod 5; c, Left uropod. Lateral plates stippled. Scale = 1.0 mm.

the Dominican Republic (type-locality), and off the Costa Rica-Panama border,
from the type-series only.

Dactylokepon sulcipes, new species
Figs. 2—5

Material examined.—Infesting Callidactylus asper Stimpson. Eastern Gulf of
Mexico. MAFLA station III-G (141); 30°01'30"N, 85°54'54”W; diver-collected;
30 m; 6 June 1974; T. S. Hopkins collector; D. L. Adkison determined host; 1 2
(gravid, holotype) USNM 172444, 1 6 (paratype) USNM 172445, host present; 2
2 (gravid), 2 6 USNM 172446 host present. BLM station 44-330701-1; 28°29/N,
84°20'W; capetown dredge; 42 m; 13 July 1976; T. S. Hopkins collector; D. L.
Adkison determined host; 1 2 (non-gravid, no d) USNM 172447 host present.
SAM station 337-2; 30°07’N, 86°45’W; trawl; 35 m; March 1977; R. L. Shipp
collector; D. L. Adkison determined host; | 2 (Guvenile), 1 dé USNM 172448 host
present.

Description.—Female (Figs. 2—4): Length 4.6-5.7 mm; width across pereo-
meres 3 or 4, 2.1-2.9 mm. Distortion angle less than 10°.

Head: frontal lamina prominent, anterior border scalloped, with a few protu-
berances on dorsal surface. Eyes small, at posterior margin of frontal lamina and
lateral edge of cephalogaster, visible in lateral view. Antenna | of 3 segments;
basal segment with 2 setae; second segment, with 5 setae Guvenile with 5 setae);
distal segment, 9 setae as terminal tuft Guvenile with 5 setae in tuft. Antenna 2
of 4 segments; basal segment lacking setae; second segment, with 5 setae Guvenile
with 3 setae); third segment with 5 setae (juvenile with 4 setae); distal segment

704 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Dactylokepon sulcipes, female, holotype: a, Dorsal view; b, Right lateral plate and pleopod
1; c, Right lateral plate and pleopod 2; d, Right lateral plate and pleopod 3; e, Right lateral plate and
pleopod 4; f, Right lateral plate and pleopod 5; g, Uropods. Lateral plates stippled. Scale = 2.0 mm
for fig. a.

with 5—7 setae (juvenile with 5 setae). Maxilliped rectangular, palp covered with
spines on medial and anterior borders. Posterior lamina with 2 pairs of projec-
tions; median pair of projections, small medially covered hook; lateral pair of
projections, either simple or multilobed, larger than median pair.
Pereon: pereopods increasing in length posteriorly. Dorsal bosses present on
pereomeres I-4; coxal plates present on all pereomeres; tergal area greatly de-
veloped on pereomeres 2-4, with tubercles present on tergal area and along suture
line between tergal area and dorsal bosses; a few tubercles present on dorsal
bosses. Tubercles on lateral margin of pereomere 7 appear to interdigitate with
setae on posterior edge of oostegite 5. Exterior surface of oostegite 5 with pos-

VOLUME 95, NUMBER 4 705

Fig. 3. Dactylokepon sulcipes, female: a, Pleon, ventral view showing trough for pereopod 7 in
oostegite 5, right side; b, Dorsal view of pereomeres 4—7; c, Maxilliped with spur; d, Posterior lamina
(spur shown in c removed); e, Posterior lamina; f, Antennae; g, Oostegite 1 and pereopod 1, exterior
view; h, Internal ridge of g; i, Internal ridge; j, Internal ridge. Figures from holotype, a—d, g, and h.
Figures from paratype female USNM 172447, f and i. Figure from USNM 172446, e.

706 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Dactylokepon sulcipes, immature female, USNM 172448: a, Dorsal view; b, Left antennae;
c, Left oostegite 1, exterior view. Scale = 1.0 mm for fig. a.

terolateral area with crenulate ridge arising at posterior edge and continuing an-
teriorly to near base of pereopod 6; second smaller ridge of protuberances present
anterior to larger ridge. Oostegite | anteriorly rounded with irregular edge; in-
ternal ridge digitate, one digitate lobe visible in dorsal view; posterior plate with
posterior edge digitate, posterolateral point present.

Pleon: six segments narrowing posteriorly. Lateral plates long, digitate, de-
creasing in length posteriorly. Pleopods 5 biramous, pairs decreasing in length
posteriorly; exopod subequal in length to lateral plate. Endopod of pleopod |
nearly % length of exopod, ratio of endopod to exopod decreasing posteriorly
until pleopod 5, when endopod 4 length of exopod. Uropod digitate, uniramous,
nearly as long as lateral plate on pleomere |.

VOLUME 95, NUMBER 4 707

Fig. 5. Dactylokepon sulcipes, male: a, Dorsal view, elongate form; b, Pleon of a, ventral view;
c, Dorsal view, compact form; d, Pleon of c, ventral view; e, Left antennae; f, Pereopod 1; g, Pereopod
2; h, Pereopod 4; i, Pereopod 7. Figures from USNM 172445, a, b, and e-i. Figures from USNM
172446, c and d. Scale = 1.0 mm for figs. a and c.

Male: (Fig. 5). Two forms found, first form elongate (Fig. 5a), other form
compact (Fig. 5c). Length 1.2-2.4 mm; width across pereomere 2 or 3, 0.50—
0.56 mm.

Head: narrower than pereomere |. Eyes present at posterolateral margin. First
antenna of 3 segments; basal segment with 6 subapical setae; second segment
with 9 setae in subapical ring; distal segment, 8-11 setae as terminal tuft. Second
antenna of 5 segments; basal segment, lacking setae; second segment with | seta;
third segment with 3 apical setae; fourth segment with 3-5 apical setae; distal
segment with 10—13 setae as terminal tuft. Maxillipeds not seen.

Pereon: of seven segments; pereomere | width greater than width of pereo-
meres 6 or 7, pereomere 7 narrowest. Pigment spots generally present on dorsal
surface of pereomeres. Legs decreasing in length posteriorly, pereopods | and 2
with dactyli and propodi enlarged, other pereopods normal. Midventral tubercles
present on all pereomeres.

Pleon: of six segments; pleomere | distinctly narrower than pereomere 7. Mid-

708 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ventral tubercle present on pleomere 1, often present on pleomere 2. Pleopods
uniramous, tuberculate, 5 pairs decreasing in size posteriorly. Uropods absent.
Pleomere 6 Y-shaped, setae present on both posterior and lateral borders; anal
cone present though generally indistinct.

Distribution.—Known only from the type-series collected in the eastern Gulf
of Mexico.

Etymology.—The specific name from Latin is a combination of sulcus (groove)
+ pes (foot) alluding to the groove in the fifth oostegite into which the seventh
pereopod appears to fit.

Discussion.—Dactylokepon sulcipes is most similar to D. caribaeus. Both
species are found on members of the Leucosiidae, D. caribaeus is found on
Iliacantha subglobosa and I. liodactyla, while D. sulcipes is found on Callidac-
tylus asper. The female of D. caribaeus differs from the female of D. sulcipes in
that the first oostegite has different ornamention on the internal ridge, there is no
posterior lateral point, the pereon has fewer tubercles, oostegite 5 is without
ridges, and the endopods of pleopods are longer. The lack of midventral tubercles
on the pereon and pleon differentiates the male of D. caribaeus from D. sulcipes.

The two forms of males found are thought to be artifacts of fixation. The
elongate form differs from the compact form by the medial separation of its
pereomeres; in the compact form the pereomeres are in contact except for the
lateral margin.

Of the five parasitized Callidactylus asper, one was a male, two were mature
females, and two were immature females. Four female bopyrids were found in
the left branchial chamber of the host while one female was found in the right
branchial chamber.

Acknowledgments

This paper represents part of a thesis submitted to the Biology Department,
University of West Florida, as partial fulfillment of a Master of Science degree.
I am indebted to Dr. T. S. Hopkins of the Dauphin Island Sea Lab, Dauphin
Island, Alabama for providing specimens, financial support, working space, and
the opportunity to make several trips to the Smithsonian Institution. Dr. R. L.
Shipp, University of South Alabama, Mobile, provided material. Material col-
lected by T. S. Hopkins was collected with support of the Bureau of Land Man-
agement contracts to the Florida State University System Institute of Oceanog-
raphy numbers 0855-CT4-11 and 0880-CT5-30 via subcontracts to him. I am grateful
to Dr. Richard W. Heard and Dr. Brian F. Kensley for answering queries and
sharing ideas during this study. I am grateful to Dr. A. E. Smalley for reading
and making suggestions on this manuscript. The type-series is deposited in the
Smithsonian Institution (USNM).

Literature Cited

Markham, J. C. 1975. New records of two species of parasitic isopods of the bopyrid subfamily
Ioninae in the western Atlantic.—Crustaceana 29:55—67.

Biology Department, University of West Florida Pensacola, Florida 32504.
Present Address: Biology Department, Tulane University, New Orleans, Loui-
siana 70118.

PROC. BIOL. SOC. WASH.
95(4), 1982, pp. 709-713

NOTALPHEUS IMARPE: A NEW GENUS AND SPECIES
OF SNAPPING SHRIMP FROM WESTERN SOUTH
AMERICA (DECAPODA: ALPHEIDAE)

Matilde Méndez G. and Mary K. Wicksten

Abstract.—Notalpheus imarpe is described from off Paita, Peru. The new ge-
nus resembles Nennalpheus Banner and Banner, but differs in the rostrum, major
chela, and second pereopod.

Recent collecting off the coast of Peru has resulted in the finding of several
new species of caridean shrimps. A most unusual find is a deep-water snapping
shrimp (family Alpheidae) that does not fit into any known genus. The new genus
and species is described herein.

Notalpheus, new genus

Diagnosis.—General body form as usual for members of the family Alpheidae.
Rostrum triangular, acute and short, with dorsal carina. Without orbital hoods.
Pterygostomial margin rounded, not protruding. Eyes well developed, visible at
least in lateral view, partly concealed in dorsal view. Orbitorostral process lack-
ing.

Antennular peduncle relatively large, stylocerite with lateral tooth well devel-
oped. Scaphocerite normal with tooth and squamous portion well developed.
Carpocerite long, basicerite with inferolateral margin pointed.

Mouthparts similar to those in Alpheus.

First cheliped with chela enlarged and carried extended; chela proper carried
in inverted position. Fixed finger bearing rounded teeth of various sizes; dactyl
without teeth, compressed. Carpus quadrangular, without teeth.

Carpus of second pereopod with 5 articles, the first not longer:than the sum of
the other 4.

Third pereopod slender, with ischium bearing 2 small spines; propodus, carpus
and merus with few long setae; dactyl unknown.

Second pleopod of male bearing appendix masculina. First 4 pleura of abdom-
inal segments rounded; fifth with posterior margin slightly projecting and sub-
acute. Posterior margin of sixth abdominal segment lateroventrally projecting into
triangular articulated pleuron.

Telson slender, with dorsal and posterolateral spines well developed. Uropods
normal.

Branchial formula as in Alpheus: 5 pleurobranchs, | arthrobranch, 8 epipodites
with mastigobranchs on bases from third maxilliped to fourth pereopod, seto-
branchs from first to fifth pereopods.

Notalpheus imarpe, new species
Figs. 1, 2

Material.—Holotype, male, total length 27.9 mm; off Paita, Peru (5°03’S,
81°20'W), 143 m, stomach contents of sciaenid fish (Larimus pacificus); R/V

710 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

0.5 cm

- Me

=~ ie ALA
USE SSS B

crane

Sn

LP»
Wp
Ds

Wj re
WIN IN4,

1 NN
ns \l .
Q ee

Fig. 1. Notalpheus imarpe: A, Lateral view of male from off Paita, Peru; B, Third maxilliped,
lateral view; C, Anterior region, dorsal view; D, Lateral view with carapace partly removed, showing

gill structure; E, Rostrum, enlarged.

VOLUME 95, NUMBER 4 Wl

Humboldt cruise 8103-04, Lance 15, 19 March 1981, Walter Elliott, collector;
Instituto del Mar del Peru.

Description.—Specimen somewhat damaged. Anterior part of cephalothorax
detached, right cheliped and dactyls of third-fifth pereopods missing.

Rostrum triangular, short and sharp, with dorsal carina. No orbital hoods. Flat
orbital teeth present. Anterior margin of carapace gradually rounded, without
teeth. Cornea visible dorsally and laterally. Second antennular article 1.4x as
long as visible part of first, 1.6x length of third article. Stylocerite reaching nearly
to end of first antennular article. Scaphocerite with external margin straight, with
lateral tooth somewhat longer than squamous part, exceeding half of length of
third antennular article. Carpocerite exceeding scaphocerite.

Mouthparts as usual in the family. Ratio of articles of third maxilliped beginning
with base 10:3:7.

Right cheliped missing. Left cheliped with chela carried in inverted position.
Chela long and thin, of generally subtriangular form, laterally compressed, length
4.04.2 breadth, fingers occupying distal 0.54. Palm suboval. Fixed finger more
compressed than palm, having on proximal two-thirds of its length series of 11-
12 rounded teeth of irregular size and other smaller inconspicuous ones, with
obvious margin along length of edge. Dactyl very compressed, laminate, without
teeth, with obvious border along length of its edge. Both fingers ending in points,
crossing each other. Carpus quadrangular, with distal margins slightly produced

dorsally and rounded. Merus 3x as long as wide, with superior and inferior
margins rounded.

Second pereopods with 5 carpal articles, with ratio of 10:3.6:3.6:3.6:5.1.

Third pereopod more or less slender. Propodus, carpus, and merus with few
long setae, ischium with 2 spines on inferior margin. Fourth and fifth pereopods
similar. Dactyls of these appendages missing.

Posterior margin of sixth abdominal segment lateroventrally projecting into
triangular articulated pleuron. Telson slender, with 2 pairs lateral spines.

Discussion.—When we first examined this shrimp, we thought that it might be
a species of Nennalpheus Banner and Banner, 1981. Like members of this genus,
our shrimp carries its chelae extended, not folded back. The general shape of the
chelae and frontal region are similar. However, species of Nennalpheus lack a
rostral carina. The fingers of the major chelae bear rounded and ‘‘exactly fitting’
teeth in the proximal half. The carpus in lateral view has a “‘rounded subrectan-
gular shape.’ The distal margins of the carpus extend into acute or rounded flat
teeth. The ratio of the carpal articles of the second pereopod is 10:2.1:2.1:5:2.2.
Our shrimp, however, has a rostral carina. Only the fixed finger of the chela bears
teeth. The carpus of the cheliped is quadrangular, without teeth. The ratio of
articles in the carpus of the second pereopod is 10:3.6:3.6:3.6:5.1.

Notalpheus, like Nennalpheus, contains shrimps from the continental shelf.
The two species of Nennalpheus, N. sibogae (De Man) and N. inarticulatus
Banner and Banner, live at depths of 70-222 m. The former has been collected
in Indonesian waters, the latter lives off the Philippines (Banner and Banner 1981).
Our new species was collected from the stomach of a fish caught at 143 m. Like
other alpheid shrimps, it may burrow or live in cracks.

Notalpheus imarpe is the eleventh species of alpheid shrimp to be reported off

712 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ss

Fig. 2. Notalpheus imarpe: A, Posterior region of abdomen showing articulated sixth abdominal
pleuron; B, Cutting edge of chela, enlarged; C, Lateral view of cheliped; D, Medial view of cheliped;
E, Male second pleopod; F, Antennular flagella; G, Carpus and chela of second pereopod; H, fourth
pereopod.

VOLUME 95, NUMBER 4 als

temperate western South America. Automate dolichognatha De Man and AI-
pheus sulcatus Kingsley have been collected off northern Peru, an area consid-
ered to be the mixture zone between the temperate Peruvian biogeographic prov-
ince and the tropical Panamic province. Holthuis (1952) reported that a specimen
of Alpheus dentipes Guérin supposedly collected at Portland Bay, western Pat-
agonia, probably was mislabelled. Athanas nitescens (Leach), normally a resident
of the eastern Atlantic, has been reported from the vicinity of Callao. This record
may be in error (Méndez 1981). Six other species are residents of the area: Be-
taeus truncatus Dana, B. emarginatus (H. Milne-Edwards), Synalpheus spini-
frons H. Milne-Edwards, Alpheopsis chilensis Coutiere, Alpheus chilensis Cou-
tiere, and Alpheus inca Wicksten and Méndez.

Etymology.—The generic name is derived from the Greek word ‘‘notos,”’
meaning southern, and Alpheus, the name of a common snapping shrimp. The
species name is an acronym of the Instituto del Mar del Peru, in honor of which
we dedicate this new species.

Acknowledgments

We thank the staff of the Instituto del Mar del Pert for use of their facilities,
biologist Walter Elliott for collecting the new shrimp, and the Consejo Nacional
de Ciencia y Tecnologia for assistance with costs of publication.

Literature Cited

Banner, A. H., and D. M. Banner. 1981. Decapod Crustacea, Alpheidae. Jn: Résultats des cam-
pagnes MUSORSTOM. I. Philippines (18-28 Mars 1976).—Collection Mémoires ORSTOM no.
91, 1(9):217-235. O.R.S.T.O.M., Muséum National d’Histoire Naturelle, Paris.

Holthuis, L. B. 1952. Reports of the Lund University Chile Expedition 1948-49. 5. The Crustacea
Decapod Macrura of Chile-—Lunds Universitets Arsskrift. N.F. Avd. 2, Bd, 47, nr 10: 1-109.

Méndez, M. 1981. Claves de identificacion y distribucion de los langostinos y camarones (Crustacea:
Decapoda) del mar y rios de la costa del Peru.—Boletin del Instituto del Mar del Peru 5:1—
170.

(MMG) Area de Invertebrados, Instituto del Mar del Peru, Apartado 22, Callao,
Peru; (MKW) Department of Biology, Texas A&M University, College Station,
Texas 77843-3258.

PROC. BIOL. SOC. WASH.
95(4), 1982, pp. 714-747

SOUTH AMERICAN FRESHWATER NEEDLEFISHES OF THE
GENUS POTAMORRHAPHIS (BELONIFORMES: |
BELONIDAE)

Bruce B. Collette

Abstract.—Potamorrhaphis and Belonion share several specialized characters:
round caudal fin; few large pharyngeal teeth; pectoral-fin rays reduced to eight
or fewer; elongate nasal barbel; and an association of the expanded first neural
spine with the supraoccipital crests and exoccipital flange which is unique to these
two genera. Potamorrhaphis has a long caudal section of its body and many
caudal vertebrae and dorsal- and anal-fin rays. Three species are recognized: P.
eigenmanni Miranda Ribeiro, P. guianensis (Schomburgk), and P. petersi Col-
lette. They show increasing numbers of dorsal- and anal-fin rays, vertebrae, and
predorsal scales from P. eigenmanni to P. guianensis to P. petersi. They form a
similar series from south to north with P. eigenmanni concentrated in the Para-
guay-Parana system, P. guianensis throughout the Amazon and the Guianas, and
P. petersi in the upper Orinoco. Apparently, P. eigenmanni invaded the Amazon
through the Mato Grosso into the Upper Rio Madeira in Bolivia and Brazil.
Similarly, P. petersi seems to have moved through the Rio Casiquiare into the
upper Rio Negro. Geographic variation in meristic and morphometric characters
was studied in the three species. Four populations of P. petersi differ only in
mean number of total vertebrae. There is variation in meristic characters within
and between major populations of P. guianensis (Orinoco, Guianas, and Ama-
zon). Generally, counts are low in the Orinoco, intermediate in the Guianas, and
high in the Amazon. These three populations differ significantly in P,—P, and P.—
C distances, pelvic fin length, interorbital width, and head width. In P. eigen-
manni, there is a general pattern of decreasing counts from the Mamoré, to the
Guaporé and Beni (all tributaries of the upper Madeira) and finally to the Para-
guay. The means for all seven meristic characters for the combined Madeira
populations are significantly different statistically from those in the Paraguay.
The Paraguay and Madeira populations also differ significantly in seven of 12
morphometric characters. Recognition of P. eigenmanni confirms the number of
species in the family Belonidae as 32 and the known neotropical species as nine
in four genera: Potamorrhaphis (3), Belonion (2), Pseudotylosurus (2), and Stron-
gylura (2).

This paper completes the review of the genus Potamorrhaphis begun with the
description of P. petersi (Collette, 1974a). A third species in the genus, P. ei-
genmanni, is validated herein from the Paraguay-Parana river system and the
upper Rio Madeira in Bolivia and Brazil.

Methods and Materials

The methods used are similar to those described in previous papers in this
series (Collette 1966, Collette 1974a—-c). Body length (BL) is substituted for stan-

VOLUME 95, NUMBER 4 TAS

—=

43}

P. petersi
s ®
S37 & :
=
rn P. guianensis < \.
SS STRONGYLURA < ABLENNES

< a |
S < i.
< ‘
UO

3] P. eigenmanni ‘ ae

~ TYLOSURUS

BELONE
2S) 7

a 86 98
TOTAL VERTEBRAE

Fig. 1. Relationship of caudal to total vertebrae in the 3 species of Potamorrhaphis and the other
genera of Belonidae.

dard length because many specimens have broken beaks and because upper jaw
length is markedly allometric in some needlefishes. BL is defined as the distance
from the posterior margin of the opercular membrane to the caudal base. Ver-
tebral counts were made from radiographs, and counts of dorsal- and anal-fin
rays made from the specimens were checked against the radiographs. Analyses
of variance (ANOVA) were conducted on the frequency distributions for each of
seven meristic characters to test for intraspecific differences. When an ANOVA
was significant (P < 0.05), a Student Newman-Keuls Multiple Range Test (SNK)
was performed to ascertain which means differed significantly from others. Twelve
measurements (listed in Table 12) were made in addition to body length. Analysis
of covariance (ANCOVA) was performed on regressions of body parts on body
length to test for intraspecific and interspecific differences. If the F value for
regression was significant at the 0.05 level, the F value for slopes was examined.

716 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

‘Ss

Fig. 2. Potamorrhaphis guianensis: a, Lateral line canals on anterior part of body; b, Pharyngeal
dentition; c, Nasal fossa and nasal barbel; d, Interorbital canal; e, Preorbital bone and canal.

If this was significant, the Newman-Keuls Multiple Range Test was performed
(Q value) to see which means differed significantly from others. If the F value
for regressions was significant and the F value for slopes was not significant, the
F value for intercepts was examined. If this was significant, the SNK was per-
formed as for the slopes.

Abbreviations used for the institutions cited herein are as follows: AMNH—
American Museum of Natural History, New York; ANSP—Academy of Natural
Sciences, Philadelphia; BMNH—British Museum (Natural History), London;
CAS—California Academy of Sciences, San Francisco; FMNH—Field Museum

VOLUME 95, NUMBER 4 717

Fig. 3. Caudal skeleton of Potamorrhaphis guianensis.

of Natural History, Chicago; INDERENA—Bogota, Colombia; IRSNB—Institut
Royal des Sciences Naturelles de Belgique, Brussels; [U—Indiana University,
collections now at CAS, designated CAS-IU; MACN—Museo Argentina Ciencias
Naturales, Buenos Aires; MBUCV—Museo de Biologia, Universidad Central de
Venezuela, Caracas; MCZ—Museum of Comparative Zoology, Harvard Univer-
sity, Cambridge, Massachusetts; MNHN—Muséum National d’Histoire Natu-
relle, Paris; MNM—Museo Nacional, Montevideo; MNRJ—Museu Nacional, Rio
de Janeiro; MSNG—Museo Civico di Storia Naturale, Genoa; MZUSP—Museu
de Zoologia, Universidade do Sao Paulo; NHMV—Naturhistorisches Museum,
Vienna; RMNH—Rijksmuseum van Natuurlijke Historie, Leiden; SMF—Senck-
enberg Museum, Frankfurt am Main; SU—Stanford University, collections now
at CAS, designated CAS-SU; UAIC—University of Alabama Ichthyological Col-
lections, University, Alabama; UMMZ—University of Michigan Museum of Zo-
ology, Ann Arbor; USNM—National Museum of Natural History, Washington,
D.C.; ZMA—Zoological Museum, Amsterdam; ZMH—Zoological Museum,
Hamburg.

Potamorrhaphis Gunther

Belone (Potamorrhaphis) Gunther, 1866:256 (original description; type-species
B. (P.) taeniata Gunther by subsequent designation of Jordan & Fordice 1887:
359, =Belone guianensis Schomburgk).

Pomatorrhaphis.—Bleeker, 1871:43 (misspelling of Potamorrhaphis although
considered as an available name by Whitley, 1970:243).

Potomarrhaphis.—Fowler, 1914:277 (misspelling of Potamorrhaphis).

Potamorhaphis.—von lthering, 1940:621 (misspelling of Potamorrhaphis).

Potamorhamphus.—Banarescu, 1970:321 (misspelling of Potamorrhaphis).

718 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Se ae SONS

+S ss

Fig. 4. Three species of Potamorrhaphis: a, P. petersi, Upper Rio Orinoco, 134 mm BL, AMNH
9619; b, P. guianensis, Rio Meta, Orinoco, 119 mm BL, ANSP 116533; c, P. guianensis, Rio Urubu,
Amazon, 108 mm BL, USNM 179527; d, P. eigenmanni, Rio Paraguay, 123 mm BL, UMMZ 207901.

Potamoraphis.—Damon, 1971:8 (misspelling of Potamorrhaphis).
Potamorraphis.—Stone, 1976:24—25 (misspelling of Potamorrhaphis).
Potamorrhamphis.—Travers, 1981:857, 863 (misspelling of Potamorrhaphis).

A number of papers referred only to Potamorrhaphis, presumably on the pre-
sumption that the genus was monotypic and contained only P. guianensis. These
are: Regan 1911:331 (74 vertebrae), 332 (in key, pharyngeal dentition); Nichols
and Breder 1928:439 (Potamorrhaphis a specialized freshwater offshoot of Stron-
gylura with a convex caudal fin); Fernandez Yépez 1948:142 (position of dorsal
fin relative to anal fin), 143 (figure), 144 (in key); Collette and Berry 1965:389
(generic characters); Géry 1969:843 (range), 845 (fig. 5); Astakhov 1980:191 (among
freshwater genera), 192 (reference to Collette 1966).

Diagnosis.—Posterior part of body elongate compared to anterior, 28-44 caudal
vertebrae constituting 43-52% of 64-85 total vertebrae (compared to 34-42% in
other Belonidae, Fig. 1). Dorsal and anal fins very long, with 27-43 and 24-39
rays respectively. Dorsal-fin rays more numerous than in any other needlefish
(except for slight overlap with Tylosurus acus melanotus, 24-27). Body lateral
line with both dorsally and ventrally directed short secondary tubes (Fig. 2a).
Pseudotylosurus also has dorsal and ventral secondary canals (Collette 1974b:
fig. 2) but these are much longer than in Potamorrhaphis. Other belonids have
the secondary canals directed ventrally (Parin and Astakhov 1982:fig. 12). Pecto-
ral-fin rays usually 7-8 (5-6 in Belonion; 8—11,.usually 9-10 in Pseudotylosurus ;
9-15 in other belonid genera). Principal caudal-fin rays reduced from usual 7+8
to 5+6, branched rays reduced from 6+7 to 4+5 (Fig. 3).

Other differentiating characters include: anterior projection of the flattened first
neural spine fitting between prominent flanges of the exoccipital and the paired

VOLUME 95, NUMBER 4

Table 1.—Numbers of dorsal-fin rays in populations of the 3 species of Potamorrhaphis.

Species Ay A B®)

P. petersi
Tomo
Upper Orinoco
Casiquiare
Negro

Species total

P. guianensis
Orinoco

Lower O.
Middle O.
Meta

Total Orinoco

Guianas

Kaituma
Essequibo
Demarara
Nickerie
Saramacca
Suriname
Maroni
Mana

Total Guianas

Amazon
Solimoes 4
Negro
Middle A.
Lower A 1
Xingu
Araguia
Tocantins
Para

Total Amazon
Species total

P. eigenmanni

Madeira
Madeira
Mamoré
Beni Oy
Guaporé 2 Ta
Total Madeira 8 13
Paraguay Ded S32
Species total 2 WS AS

30

—

ila
73)

31

—

13

20

31
3p)

32

15

a)

139

Rl] WH le

33

QO | — —

Sl

— N
Wee HRhNNNH NH

Nn
oO

34

—
NN Bh S|] WN

ill
36
58

35

15
19

36 37 «438

QQ pee

39

40 41 42 43

x

333)

311.1
28.7
30.2

30.0
29.0
29.4

719

10
15
33

Sy)
66
ZS)

supraoccipital crests as in Belonion; 3 pairs of upper pharyngeal tooth plates (Fig.
2b) as in most other Belonidae (except Belonion and Xenentodon) but the third
pair (UP;) greatly reduced; lower pharyngeal bone (fused fifth ceratobranchials)
triangular (Fig. 2b), not elongate as in Pseuwdotylosurus and Xenentodon; nasal

720 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Numbers of anal-fin rays in populations of the 3 species of Potamorrhaphis.

Species DAD DSN 26). Ta 28" D9) ABO 48, BO Bp aA Soper Sie Ses Cie 3 Oo nomae n
P. petersi
Tomo 1 2 I
Upper Orinoco 2 ch a SO De O8 6
Casiquiare 1 34 1
Negro |e res Aes) 36 8
Species total 1 = 4 2° 3 -3 2. “‘Aas35.8eelG

P. guianensis

Orinoco
Lower O. Deel Oe wed foe ye
Middle O. DAIS) Si e.02 1 Way 8
Meta it 28a wee
Total Orinoco AV NG. 2a a l 27.6 48
Guianas
Kaituma 1 1 28.5 2
Essequibo 27 Ss Blo ey 28.6 44
Demarara oD 1 2 I 29.3 6
Nickerie 10 8 1 27 Sie lS
Saramacca i 1 28.5 4
Suriname 1 8 19 6 1 PAPO) B2)5)
Maroni 1 A 28.1 7
Mana 1 Di, 1
Total Guianas le 22 SO O27 to 1 28.2 118
Amazon
Solimoes i Zz 14 9 1 ea Sy]
Negro DO od (OR ET d call 30 2a 8
Middle A. Sy, 2! 28.4 9
Lower A. | Oe 2304 £424 MID eS 1 27.9 96
Xingu Sets Sues 29.0 2)
Araguia 2 29.0 2
Tocantins 2 28.0 2
Para l 10 13 10 ] 29.0 35
Total Amazon I ZO AGE Se 845 Sie Oneal 28.2 228
Species total 125) S45 Se ae 40 asl Oe 28.1 394
P. eigenmanni
Madeira
Madeira 1 28 1
Mamoré 1 — 8 DLS 10
Beni 2 Flor 3 25.1 15
Guaporé A ow 0 3 26.4 33
Total Madeira 2 15 20 10 12 MGes SS)
Paraguay Se | 5 — 1 25.4 - 65

Species total 9 46 41 15 12s 25.8 124

barbel elongate, projecting out of nasal fossa (Fig. 2c) as in Belonion and Xe-
nentodon; caudal fin rounded as in other freshwater genera; caudal peduncle
compressed, with no trace of a lateral keel; supraorbital canal simple, with 4
pores on each side (Fig. 2d), and without any secondary canals such as are present
in most other belonid genera (Parin and Astakhov 1982:fig. 4).

VOLUME 95, NUMBER 4

Table 3.—Numbers of pectoral-fin rays in the 3 species of Potamorrhaphis.

Species

P. petersi
Tomo
Upper Orinoco
Casiquiare
Negro

Species total

P. guianensis
Orinoco
Lower O.

Middle O.
Meta

Total Orinoco

Guianas

Kaituma
Essequibo
Demarara
Nickerie
Saramacca
Suriname
Maroni
Mana

Total Guianas

Amazon

Solimoes
Negro
Middle A.
Lower A.
Xingu
Araguia
Tocantins
Para

Total Amazon

Species total
P. eigenmanni
Madeira

Madeira
Mamore
Beni

Guaporé

Total Madeira
Paraguay

Species total

6

N

7

8

Da oy N
N

721

Description.—Dorsal-fin rays 27—43; anal-fin rays 24-39; pectoral-fin rays 7 or
8, rarely 6. Precaudal vertebrae 35—42; caudal vertebrae 28—44; total vertebrae
64-85. Predorsal scales 77-143. Vomerine teeth absent. Gill rakers absent. Two
gonads present, right about as long as left, left/right = 0.810-1.176, no sexual
dimorphism in ratio. No posterior lobe in dorsal fin of juveniles. Juvenile “‘half-

722 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

beak’’ stage either absent or upper jaw grows to near tip of lower jar by an early
stage (by 35 mm BL). Anterior rays of anal fin longer than posterior rays and
forming a distinct lobe. Branchiostegal rays 10-11. Preorbital canal with short,
ventrally directed branch midway along its anterior margin (Fig. 2e).

Potamorrhaphis petersi Collette
Fig. 4a

Potamorrhaphis guianensis (not of Schomburgk, 1843) Mago Leccia, 1971:9 (Rio
Casiquiare, Venezuela).

Potamorrhaphis petersi Collette, 1974a:34—38, figs. la, 2 (original description;
Upper Rio Orinoco and Rio Negro). Cala, 1977: 13 (listed, Colombian Orinoco).

Diagnosis:—Most similar to P. guianensis from which it differs most markedly
in higher numbers of dorsal- and anal-fin rays, vertebrae, and predorsal scales
(Tables 1-7).

Types.—Potamorrhaphis petersi Collette, 1974. Holotype: USNM 210546 (162
mm); Colombia, Laguna Coco NE of Puerto Inirida, pool near junction Rio Gua-
viare and Rio Inirida; 17 Jan 1972; P. Cala. D 39; A 34; P, 8-7; vertebrae 40 +
40 = 80; predorsal scales 130; P,—P. distance 73.0 mm; P,—C distance 85.3 mm.
Paratypes: 7 specimens, USNM 210547; AMNH 9619; MNHN 87-655-6 and USNM
210861; MBUCV-V-6132; and CAS 27587 (see Collette 1974a:38 and Material
Examined).

Geographic variation.—The four populations of P. petersi examined differ in
only one meristic character, total number of vertebrae. The means for the Tomo
(77.0), Casiquiare (80.0), and Rio Orinoco plus Rio Negro (82.3 and 82.8) are
significantly different. There are no significant morphometric differences between
any of the populations.

Material examined.—RIO ORINOCO: 8 specimens (97.5—162) from 5 collec-
tions. USNM 210546 (1, 162) Colombia, Laguna Coco NE of Puerto Inirida, pool
near junction of Rio Guaviare and Rio Inirida; 17 Jan 1972; P. Cala; holotype of
P. petersi. USNM 210547 (1, 137), same data as holotype. INDERENA G. A.
P-0066 (1, 111), Colombia, Rio Tomo near entrance into Rio Orinoco. AMNH
9619 (1, 133) Venezuela, Cano Pescado, 8 km N of Esmeralda; 9 Mar 1929; G.
H. Tate. MNHN 87-655-6 (2, 97.5—123) and USNM 210861 (1, 108) Venezuela,
Amazonas, Rio Atabapo at San Francisco de Atabapo; Oct 1886; J. Chaffanjon.
MBUCV-V-6132 (1, 124), Venezuela, Amazonas, Cano Beripamoni, tributary of
Rio Casiquiare; 29 Jan 1969; F. Mago Leccia, J. Mosco, A. Machado.

RIO NEGRO, Brazil: 8 specimens (64.8—157) from 4 collections. CAS 27587
(1, 92.7) Sao Gabriel above Camanos; | Feb 1925; C. Ternetz. USNM 222571 (2,
137-145) Ilha de Buiu-acu-Tapuruquara; 6 Feb 1980; M. Goulding 959. USNM
222572 (4, 84.8-157), Ilha de Tamaquaré-Tapuruquara; 11 Oct 1979; M. Goulding
55. USNM 222573 (1, 152), mouth of Rio Urubaxi; 3 Feb 1980; M. Goulding.

Potamorrhaphis guianensis (Schomburgk)
Fig. 4b—c

Belone Guianensis Schomburgk, 1843:131—132, color pl. 1 (original description;
Padauiri River, Guiana).

VOLUME 95, NUMBER 4 128

Table 4.—Numbers of precaudal vertebrae in the 3 species of Potamorrhaphis.

Species 35 36 37 38 39 40 Al 42 z n
P. petersi
Tomo | 39 1
Upper Orinoco 3 3 40.5 6
Casiquiare 1 40 1
Negro y) 3) I 40.4 8
Species total 6 1 40.3 16
P. guianensis
Orinoco
Lower O. i 2 8 7 l 39.3 19
Middle O. 2 11 10 39.3 23
Meta 1 40 1
Total Orinoco l 4 19 18 1 39.3 43
Guianas
Kaituma D 39.0 D)
Essequibo He 7 2 38.5 21
Demarara 2 39.0 y,
Nickerie 4 7 4 38.0 IS
Suriname 1 5) 9 39.3 25
Maroni 1 38 1
Total Guianas 4 ” | 30 il 38.7 66
Amazon
Solimoes 4 7 4 4 38.4 19
Negro 3 7 4 40.1 14
Middle A. D 1 38.3 3
Lower A. 10 10 14 W 38.4 41
Xingu 3 6 1 38.8 10
Araguia 2 39.0 2
Tocantins 1 l 38.5 D,
Para 1 16 il th 39.4 28
Total Amazon 14 24 47 30 38.9 119
Species total 19 49 96 59 5 38.9 228
P. eigenmanni
Madeira
Mamoré 4 5 1 Sol 10
Beni 1 9 4 S72 14
Guaporé 3 17 10 37/3 32
Total Madeira 4 30 19 37.4 56
Paraguay D 28 il 2 36.4 58)
Species total 2 32 Sil 21 - 3 36.9 109

Belone scolopacina Valenciennes in Cuvier and Valenciennes, 1846:428—429
(original description; La Mana, Cayenne, French Guiana).

Tylosurus guianensis.—Miuller and Troschel, 1848:626 (after Schomburgk).

Belone (Potamorrhaphis) taeniata Gunther, 1866:256 (original description; Bra-
zil).—Couto de Magalhaes, 1931:149-150 (description, Amazon), fig. 76.—
MacDonagh, 1938:189 (comparison of types with 8 P. eigenmanni from the
Paraguay River by Dr. Trewavas, BMNH).

724

Table 5—Numbers of caudal vertebrae in the 3 species of Potamorrhaphis.

Species 28229" 30) 3 32

P. petersi
Tomo
Upper Orinoco
Casiquiare
Negro

Species total

P. guianensis

Orinoco
Lower O. |
Middle O. 6
Meta
Total Orinoco lh Ale
Guianas
Kaituma
Essequibo
Demarara
Nickerie oranges,
Suriname 1
Maroni
Total Guianas 3 al
Amazon
Solimoes t 10
Negro
Middle A.
Lower A. 1 Pal
Xingu
Araguia
Tocantins
Para
Total Amazon [ene
Species total 1” 6 30
P. eigenmanni
Madeira
Mamoré ag tS
Beni Ss ral
Guaporé oe @- stil» Ih
Total Madeira ets: Riss
Paraguay Pe 23 I ©
Species total I 3 WE Wil

33

20

20
60

CoO ;N = NM

34

26
60

35

36

37

10
10

38

39 40 41
3
1

BVA

42

43 44

30.3
30.8

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

a2
106

Potamorrhaphis taeniata.—Steindachner, 1876:96 (taken by Thayer Expedition

at several Amazonian localities).

Potamorrhaphis guianensis.—Jordan, 1887:530 (reference to type-specimen of
Belone scolopacina).—Jordan and Fordice, 1887:359-360 (synonymy, descrip-
tion, specimens from Itaituba, Brazil) —Eigenmann and Eigenmann, 1891:66
(synonymy).—A. de Miranda Ribeiro, 1915:I, 11-13 (description, 8 specimens

VOLUME 95, NUMBER 4 25

from Rio Negro at Manaus). Fowler, 1919:6 (Peruvian Amazon and Rupununi
River; P. eigenmanni considered a synonym).—Schultze, 1939: 101 (specimen
in Munich Aquarium from Georgetown, British Guiana).—Fowler, 1940a:278
(synonymy; Ucayali River basin, Peru.—-Eigenmann and Allen, 1942:46 (lower
Maranon), 48 (lower Ucayali) 58, in part (upper Amazon, Brazilian Amazon,
Guianas), 381-382 (snyonymy, distribution).—Martin, 1954:3 (Venezuela, char-
acters), 6 (in key).—Ringuelet and Aramburu, 1961:53 (listed. Mees, 1962:4
(recognized as separate monotypic genus).—Mees, 1964:314—-315 (synonymy,
description, range, P. eigenmanni considered a junior synonym).—Collette,
1966 (comparison with Belonion), tables 1-3 (numbers of vertebrae, pectoral
fin rays, and branchiostegal rays), fig. 2A (pharyngeal teeth), fig. 6B (caudal
skeleton), fig. 7B (interorbital canals)—Ovchynnyk, 1967:40 (Ecuador; distri-
bution).—Ovchynnyk, 1968:260 (Ecuador, distribution).—Cressey and Col-
lette, 1970:400 (presence of parasitic copepod, Ergasilus orientalis on Brazilian
specimens), 416 (fig. 180, distribution).—Berghegger, 1970: 199-203 (article on
keeping P. guianensis in aquaria illustrated with photographs of juvenile Xe-
nentodon cancila and with photographs of adult Xenentodon labelled as P.
guianensis, see Foster 1973:84).—Mago Leccia, 1970:89 (listed, Venezuela).—
Buen, 1972:160 (distribution, in part; photograph).—Hinegardner and Rosen,
1972:639 (DNA content; 24 haploid chromosomes).—Collette, 1974a (compar-
ison of P. petersi with populations of P. guianensis from the Orinoco, Guianas,
and lower Amazon), 37 (fig. 3, distribution of northern populations).—Kusaka,
1974:110 (fig. 161, description of urohyal of specimen from Guiana).—Bohlke
and McCosker 1975:10 (Rio Tocantins near Tucurui).—Cala, 1977:13 (Rio Man-
acacias, tributary of R. Meta, Colombia).—Hubbs and Wisner, 1980:549 (ref-
erence to pharyngeal bones in Collette 1966).—Greenwood and Lauder, 1981:
221 (protractor pectoralis muscle present).—Parin and Astakhov, 1982:278 (ref-
erence to lateralis system from Collette 1966).

Potomarrhaphis [sic] guianensis.—Fowler, 1914:277 (Rupununi R., British
Guiana).

Potamorhaphis [sic] guianensis.—von lhering, 1940:621 (Brazilian common name
pira-pucu).—Sterba, 1962:605 (description).

Tylosurus scolopacina.—Puyo, 1949:163 (description, in part, not reaching 768
mm in length; French Guiana).

Potamoraphis [sic] guianensis.—Damon, 1971:9 (description of aquarium speci-
mens).

Potamorraphis [sic] guianensis.—Stone, 1976:24—25 (photos of aquarium speci-
mens).

Potamorrhamphis [sic] guianensis.—Travers, 1981:857, 863 (interarcual cartilage
absent).

Potamorrhaphis sp. Fyhn et al., 1979: fig. 9, table 9, 60 (electrophoretic pattern
of hemoglobin, 2 Amazonian specimens).

Misidentification.—Schultz (1949:74) reported Potamorrhaphis guianensis from
the Rio Apure in Venezuela. The specimen (UMMZ 146392) was reexamined and
found to be Pseudotylosurus microps (Gunther).

Diagnosis.—Intermediate in dorsal- and anal-fin rays, vertebrae, and predorsal
scales between P. petersi, which has higher counts, and P. eigenmanni, which
has lower counts (Tables I-7).

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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VOLUME 95, NUMBER 4

728 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 7.—Numbers of predorsal scales in populations of the 3 species of Potamorrhaphis (x based
on original frequency distributions).

Species WW WD Bi 83 Dd wy bY Bl 93 95 97 499 101 103 105 107

P. petersi

P. guianensis
Orinoco

Lower O. = 1 3 i es
Middle O. 1
Meta

Total Orinoco 2 Gh Se Atay eas 1 8

&
nN
ie)
N
—
ie)

Guianas
Guyana 1 —
Nickerie 2 I 2
Saramacca-Suriname
French Guiana —

Total Guianas | 4 1 Digy 8) 4

| |
Nv
| aw |
a
iN
Nv
20

Oo

=|
n |—
a | |
o>

Amazon
Solimoes 3
Negro 1 —
Lower Amazon D
Para

Oe ctoe | a
Nn HK nN
—= ~] — VY

a |
On| Wl —
—
ON
—
ee

1
Total Amazon 3 4
Sy 10

=
L
—
N
GN
—
—

Species total Pp)
P. eigenmanni
Madeira
Madeira
Mamoré
Beni el
Guaporé

Total Madeira al
Paraguay DT Th tg BS Dee

rete Rey etal
S |
NIN Ns] —
Ccojme N/|BR KS LV
—
—= |W CO |Un = bO
o;mTN a|]s

=

i)

—

Species total 2. Eli emSi 3), 4

Types.—Belone Guianensis Schomburgk, 1843. No types known to be extant.
Original description based on a specimen taken in the Padauiri River, British
Guiana.

Belone scolopacina Valenciennes, 1846. Holotype MNHN 833 (123 mm); French
Guiana, Cayenne, Rio de la Mana; Leschenault and Doumerc. D 32; A 27; P,
7-7; predorsal scales about 100; P,—P. distance 59.2 mm; P,—C distance 60.2 mm.

Belone (Potamorrhaphis) taeniata Gunther, 1866. Lectotype: BMNH 1849.11.9:
59 (142 mm); Brazil, Para, Capim River; purchased from Mr. Stevens; herein
selected. D 32; A 30; P, 8-8; vertebrae 38 + 36 = 74; predorsal scales about 110;
P,—P., distance 69.1 mm; P.—C distance 69.9 mm. Paralectotypes: BMNH 1849.11.9:
60-61 (2, 130-153); same collection data.

Geographic variation.—Two sets of ANOVA comparisons were made for most
of the meristic data in P. guianensis. First, comparisons were made among spec-
imens from three sections of the Rio Orinoco, among eight rivers in the Guianas,

VOLUME 95, NUMBER 4 129

Table 7.—Continued.

109 iii ss Tiss I) TNL BS AS IS) ST 11838) IBIS) SAB) TIABT 11's) n x
ee a 2 ee 22 ll sie tS eee k Sop laa | = WE).3)
4 eS.) 1 —- — — — — — 1 21 109.4
| 23r ee lO220

| ene

> 3 3 0 — 1 —- — — — — — 1 45 105.7
4 Bee Ouene re Dri i2 4 ee 18 34 112.9

eae 220)
1 1 24 104.3

4 100.0
a Me Di OD oA eB 74 ~ 106.0
7 Pm OR) wie 3 a6 40045) 2 1 I 7 ee | SO WB)

— Pee Orly 2. 2 5 129
8 SMe Oo ed AD D8 G2) IMIS.S)
5 ma oO OD I l 39, LULA

ele 24 14 4" 6S 4a 2 169 113.4

Seeley 26 Tr 16 10 ST SD eet 288 110.3

eeaasens, 10 99.5
9 93.4

ie eaieli 30 97.4

(fe Deas 5 Sy
49 90.6

een | 99 93.9

and among eight tributaries or regions in the Amazon drainage (Tables 8—10).
Comparisons were then made among the three major populations: Orinoco,
Guianas, and Amazon (Table 11).

There is adequate material to compare P. guianensis from the lower Orinoco
with the middle Orinoco (Caicara area). There is also one specimen from much
further upstream, in the Rio Meta. The specimen from the Rio Meta had higher
counts, except for the number of pectoral-fin rays. Differences were significant
only for numbers of caudal and total vertebrae. Samples from the middle and
lower Orinoco do not differ from each other but do have significantly lower counts
than the specimen from the Rio Meta (Table 8).

Material was available from eight rivers in the Guianas. From northwest to
southeast they are the Katuma, Essequibo, and Demarara in Guyana; the Nick-
erie, Saramacca, and Suriname in Surinam, the Maroni (or Marowijne) on the
Surinam-French Guiana border; the Mana River in French Guiana. In the Guia-

730 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 8.—Comparison of means for meristic characters for 3 populations of Potamorrhaphis gui-
anensis from the Orinoco River (* indicates mean is significantly different from adjacent mean by
SNK, alpha = 0.05).

Fin Rays Vertebrae Scales
Population n Dorsal Anal Pectoral Precaudal Caudal Total Predorsal
Lower 19-24 BRS Des 7.0 3033 32.7 r 7.9 109.4
Middle 21-23 3S Pos) 7.0 39.3 33.0 72.3 102.0
Meta 1 33 28 7 40 35) a" 1

nas, specimens with the lowest counts were in the Nickerie River, sometimes
joined with the Kaituma (Table 9). Specimens with the highest counts were in
the Demarara River, sometimes joined with the Essequibo. Samples from the
Saramacca, Suriname, Maroni, and Mana were generally intermediate between
the groups with low and high counts. This pattern of meristic variation does not
seem to conform to any simple geographic pattern.

The Amazon River was divided into eight areas or major tributaries. From
upstream (west) to downstream (east), these are: Solimoes (or upper Amazon:
Colombia, Ecuador, Peru, and Brazil down to the confluence with the Rio Negro);
middle Amazon (Manaus to Gurupa); Xingu; Araguaia; Tocantins; and the Para
region. Samples with the lowest counts in the Amazon drainage were from the
Solimoes (Table 10). The highest counts were generally in the Rio Negro, some-
times joined by the Para population. The Tocantins, lower Amazon, Xingu, and
Araquaia populations were generally intermediate between the low and high counts.
As in the Guianas, this pattern of meristic variation does not seem to conform to
any simple geographic pattern, lowest and highest counts being from adjacent
areas.

The three major populations of P. guianensis show a general pattern of low
counts in the Orinoco, intermediate in the Guianas, and high in the Amazon
drainage (Table 11). Populations in the Orinoco and Guianas were not significantly
different in two characters (total vertebrae and predorsal scales) and those in the
Guianas and Amazon were not significantly different in three characters (dorsal-

Table 9.—Comparison of means for meristic characters for 8 populations of Potamorrhaphis gui-
anensis from the Guianas. (* indicates mean is significantly different from adjacent mean by SNK,
alpha = 0.05. Means with the same superscript do not differ from each other but do differ from means
with different superscripts).

Fin rays Vertebrae Scales
Population n Dorsal Anal Pectoral Precaudal Caudal Total Predorsal

Nickerie EID (BIS |. 27 50 2" (A138 OR SOs | 70.5! 92.0%
Kaituma 2 32.0 28.5 HA02 39.0? 3320Rle W2EO% —
Saramacca 0-4 32.3 28.5 7-03 — — — 104.3
Suriname 21-36 Ban 27.9 VSP 39532 33.6 | 72.8? ——
Maroni 1-7 32.6 28.1 Wes 38.0! 34.1 V2AL0: 100.0
Mana 0-1 B05 DY AV — — i" —
Essequibo 19-44 33.3 28.6 7.82 38.52 34.3 | bas te
Demarara 1-6 34.0 29.3 8.0? 39.0? 36.3 JSE0; ;

VOLUME 95, NUMBER 4 731

Table 10.—Comparison of means for meristic characters for 8 populations of Potamorrhaphis
guianensis from the Amazon. (* indicates mean is significantly different from adjacent mean by SNK,
alpha = 0.05. Means with the same superscript do not differ from each other but do differ from means
with different superscripts).

Fin rays Vertebrae Scales

Population n Dorsal Anal Pectoral Precaudal Caudal Total Predorsal

Solimoes ISS 34? DBAS TiN? (Bs 138.42 SESE WNC | 113.9
Tocantins 2 Boh 28.0 Jie Bksesr 33.5" 72.0 —
Lower 41-96 32S ZA se) } 7.3? = 38.4! 33.8) s 13s5)
Middle 2-9 Sei 28.4 TAP pr Bosak 34.5 T2D -—
Xingu S=10) 3255" 29.0 6.91 38.8? 34.2 | Woe)" —
Araguia 2 325 DOOM es Vir SSP 35.0 74.0 —

Para 26-38 SBmle 29.0 SE) 39 At 34.8 74.3 ¢* 112.6

Negro 12-18 34.9% 30.2 PA S400 36.4 76.4 112.9

and anal-fin rays and precaudal vertebrae). All three populations were signifi-
cantly different in numbers of caudal vertebrae and pectoral-fin rays.

Two sets of ANCOVA comparisons were also made for morphometric data in
P. guianensis. First, comparisons were made of two populations in the Guianas
and four populations in the Amazon drainage. Second, comparisons were made
among the three major populations: Orinoco, Guianas, and Amazon.

No statistically significant differences were found between the Guianas popu-
lations. Significant differences were found in 10 of 12 morphometric characters
(all but snout length and postorbital distance) among the four Amazon populations
(Table 12). Slopes differed for head length, orbit, interorbital width, head depth,
and head width. Slopes did not differ but intercepts did for P,-P, and P.-C dis-
tances, preopercle length, and pectoral and pelvic fin lengths. However, the New-
man-Keuls Multiple Range Test was able to select which population was signif-
icantly different for only three characters: P,—P. and P.-C distances and interorbital
width (Table 13). The Rio Negro population was significantly different from the
other three populations in all three characters.

Statistically significant differences in slopes or intercepts were found for eight
of the 12 morphometric characters among the three major populations of P. gui-
anensis: Orinoco, Guianas, and Amazon (Table 14). Slopes differed for interor-
bital width, head width, and pelvic fin length. Intercepts differed for P,-P. and
P,—C distances, preopercle length, orbit diameter and head depth. The Newman-
Keuls Multiple Range Test was able to determine which population or populations
differed significantly in all characters except preopercle length, orbit diameter,

Table 11.—Comparison of means for meristic characters for 3 populations of Potamorrhaphis
guianensis (* indicates mean is significantly different from adjacent mean by SNK, alpha = 0.05).

Fin Rays Vertebrae Scales
Population n Dorsal Anal Pectoral Precaudal Caudal Total Predorsal
Orinoco 36-48 310 27.6* We 3) oi B2A98 (ee LOSEV/iai\ee
Guianas OSM S20 |e 9 as 7.6* 3060 || a 33.40% V22 106.0
Amazone» 1IS=229° 3255 28.2 i pes 38.9 34.3* Vl Wa

732 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 12.—Analysis of covariance of regressions of morphometric data on body length in 4 popu-
lations of Potamorrhaphis guianensis from the Amazon drainage: Solimoes (S), Negro (N), Lower

Amazon (L), and Para (P). (* = significant at 0.05).

Character Pop. Y= CD N P Q
Head length S77 0r65520> 90045 98s 39 Resiresston 2.786 .045
NN 0506X = 95696-2895 “1 Slopes oll O11 23375
I WSR ae oe idk
IP Undsavk se Joi Gils 113}
Snout length S 0.453X — 3.881 .917 39 Regression DS .089
INS OL3DSI2K =F SLO sll IL
JL, 38D ae Oise ks Sh)
2 O.38EE« se AGP els 1}
P,-P, SS ODIB Xe 1700) 977 oe Reeression 4.236 .007
IN Ose 38208 989 Ge Slopes 0.104 958
lo Os IDS = ZT SO ST Sl Intercepts 8.349 <.001 S-2365
2 OSU. = O03 232 — lo
P.-C S 0.470X + 1.156 .974 56 Regression 5.382 .002
N 0.484X + 2.046 .995 13 Slopes 0.071 poHS
OFT 2X6 eso ly 2976S ole intercepts 10.675 <.001 Tees
le UAT Se i Sis) IG
Postorbit S 0.111X + 0.534 .923 56 Regression 2.314 .078
N 0O.111X + 0.040 .930 13
I WUD Se O04 SON 4
IP OMIM CSS eID Soy ONY)
Preopercle Ss) UU = Whit RB Sg INGESTION 3.076 .029
N 0.059X — 0.094 .94] 13 Slopes ihe fl 7 344
Ib) OWS Dae O27 72. Od, linieregatis 4.925 .003 —0.900
IP Oi0SEDK = OL059) cis) 20)
Orbit S 0.042X + 0.776 .913 56 Regression Deseo .038
N 0.034X + 1.523 .908 13 Slopes 3.070 .030 3.347
L 0.035X + 1.472 .844 75
POWER ae IIS IS 20
Interorbit S 0.063X — 0.934 .948 56 Regression 4.684 .003
N 0.043X + 0.935 .865 13. + Slopes Bye 22 | .024 4.171*
1 OUD CS IOI SS IS) .
1 OCI = O85 ~ S08 9 20
Head D Ss 1 OWTOx = l0ss 830 Sil Regression 11.016 <.001
IN OWoIbosb Oo 229 is Slooess 3.188 .026 3.053
kp OCG = O55) VS OZ
P 0.064X — 0.104 .962 20
Head W SS) WTS = 543) OI Sil Ikearessiioim 6.691 .0003
INF OOS8< sb O,337 53 IS slopes 4.529 005 3.383
I Woy = Oss44! Dil
IP OWoPke a OS Os 20)
Py S 0.126X — 0.420 .916 45 Regression 4.772 .003
ING ORS OX =) 028879207 als eslopes 1.790 al52
I Ok = IS se SO Iiintisineejatis 7.436 .0001 —5.498
I LIOR ze MON WIS 2D
Pik S 0.086% — 1.406 .914 52 Regression 5.568 .001
INE) OF093Xx = 17/661 2892 13. ~Slopes 2.404 .070
LE 0092 28 2878 oy Minitercepts 8.316 <.001 — 1.461
ES O07 0X6 Orr er or 20

VOLUME 95, NUMBER 4 733

Table 13.—Newman-Keuls Multiple Range Tests of significantly different slopes or intercepts for
regressions of morphometric data on body length in 4 populations of Potamorrhaphis guianensis from
the Amazon drainage. Regressions and analysis of covariance from Table 12. (* = significant at 0.05).

P,—P, distance Intercepts Q for ranks
1. Negro —3.82033 14 5.2360*
2. Lower = 2 O27) 1-3 6.6185*
3. Solimoes — 1.69968 1-2 5.1841*
4. Para —0.80284 2-4 0.0900
P.-C distance Intercepts Q for ranks
1. Solimoes 1.15566 14 WoZB Ney
2. Lower 1.86083 1-3 2.7316
3. Para 1.97143 3-4 4.7068*
4. Negro 2.04583
Interorbit Slopes Q for ranks
1. Negro 0.04279 14 4.1708*
2. Para 0.06098 1-3 4.7754*
3. Solimoes 0.06316 1-2 32513"
4. Lower 0.06404 2-4 0.8550

and head depth (Table 15). The Guianas population differed from those in the
Amazon and Orinoco in both P,—P, and P,—C distances. The Orinoco population
differed from those in the Guianas and Amazon in pelvic-fin length. All three
populations were significantly different from each other in interorbital width and
head width.

Material examined.—ORINOCO RIVER: 50 specimens (62.7—126 mm BL) from
19 collections. CAS SU 52683 (7, 99.0-112), 52684 (1, 101), 52686 (8, 88.4—113),
58815 (1, 96.4); CAS 28322 (3, 62.7-110), 28323 (1, 92.3); USNM 209303 (3, 87.8—
111), Venezuela, Cano Queribana into Orinoco at Caicara; May 1925; C. Ternetz.
ANSP 116533 (1, 119), Colombia, Rio Meta, Cano Emma at Finca El Viente S
of Matuzal; 4°08’N, 72°39’W; 18 Mar 1973; J. E. Bohlke, W. Saul, W. Smith-
Vaniz. USNM 219836 (12, 88.6—-114), Venezuela, Delta Amacuro at about km 65;
J. N. Baskin et al. 80-78. UCV uncat. (1, 87.3), Rio Orinoco, 8°36'24’N, 61°00’ W;;
J. N. Baskin 74-79. ANSP uncat. (1, 108) Rio Arature, 8°32'N, 61°00’W; 24 Feb
1978; J. N. Baskin 85-78. LACM uncat. (1, 123) same locality; J. N. Baskin 88—
78. FMNH uncat. (1, 89.7) Rio Orinoco, 8°32'N, 61°02’W, km 74; 25 Feb 1978;
J. N. Baskin 80-78. USNM 234948 (2, 92.0—126) Rio Orinoco, 8°29’N, 61°18'’W,
km 82; 22 Feb 1978; J. N. Baskin 64-78. USNM 234947 (1, 125) Rio Orinoco,
8°36'N, 61°48’W, km 117; 20 Feb 1978; J. G. Lundberg 47-78. USNM 234946 (2,
79.4—-106) Rio Arature, 8°32'’N, 60°52'06”W; 1979; E. C. Marsh 25-79. CAS uncat.
(1, 96.3), USNM 234944 (1, 86.8), and USNM 234945 (2, 78.7-86.3) Rio Orocop-
iche, 8°03’N, 63°40’W; 1979; J. N. Baskin 1, 2, and 5-79. GUIANAS: 144 spec-
imens (40.4-144 mm BL) from 61 collections. KATUMA RIVER, GUYANA.
CAS SU 51288 (2, 117-123), headwaters Katuma R.; 25 Apr 1958; A. Fletcher.
ESSEQUIBO RIVER, GUYANA: 54 (40.4—130) from 21 collections. CAS [UM
12580 (1, 84.0) lower Potaro R., Tumatumari; Eigenmann 1908 Exped. CAS IUM
12581 (1, 106) Essequibo R., Gluck I.; Eigenmann 1908 Exped. CAS IUM 12582
(1, 89.5) lower Potaro R., Potaro Landing; Eigenmann 1908 Exped. CAS IUM

734 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 14.—Analysis of covariance of regressions of morphometric data on body length in the 3
major populations of Potamorrhaphis guianensis: Orinoco (O), Guianas (G), and Amazon (A). (* =
signficant at 0.05).

Character Pop. Y= CD N F P Q
Head length OF OSORxe=. 09 ie 295i 18 Regression 0.610 545
G 0.541X + 4.301 .890 26
A 0.584X — 0.499 .899 109
Snout length OQ) O42IbS =A1LOID Qil7/ 18 Regression 0.534 588
G 0.400X + 2.342 .822 26
A 0.424X — 0.516 .887 109
P,—P, © 05 1SxXe = 223949 2971 30 ~=Regression 4.509 .012
G Ws. = Bai 235) 58 Slopes 0.014 .986
Ay 025 1S XG 2212 2981 ISO eeintencepts 9.003 .0002 6.483*
P.-C O 0.476X + 1.217 .959 30 Regression 5.644 .004
G 0.461X + 3.903.982 58 Slopes 0.257 174
EO OFA OP XS 22 1 oT = S50 eeintencepts 11.006 <.001 Sw Il2
Postorbit O 0.102X + 1.121 .864 30 Regression 1.988 .139
G 0.105X + 0.997 .941 59
ev OSWOX se O670. S24. a7
Preopercle O 0.052X + 0.968 .794 30 Regression 4.221 .016
G 0.063X + 0.020 .926 59 Slopes 1.604 .203
A 0.062X + 0.011 .911 171 ~~ Intercepts 6,2 .001 —0.386
Orbit O 0.033X + 1.430 .698 30 ~=Regression 32590 .030
G_ 0.039X + 1.020 .819 59 Slopes 0.714 419
A 0.038X + 1.169 .885 178 Intercepts 6.351 .002 —3.888
Interorbit O 0.045X + 0.635 .804 30 + Regression 13.829 <.001
G_ 0.056X — 0.440 .907 59 Slopes 8.675 .0002 4.865*
A 0.063X — 0.947 .935 178
Head D O 0.060X + 0.237 .856 30 Regression B25 .039
G 0.068X — 0.327 .963 57 Slopes 1.954 144
A 0.071X — 0.701 .928 160 Intercepts 4.535 012 =2 EZ
Head W O 0.041X + 1.314 — .833 30 Regression 16.332 <.001
G_ 0.059X — 0.029 .909 57 Slopes 11.306 <.001 37855)
A 0.067X — 0.870 .917 160
Jee IL, OF 02083Xx0-5 3:373 2568 23 Regression 2.268 .106
Ge One — 02 S716 54
IN WN2ZIPS SOU37 wets) MANS
Pek O 0.051X + 1.978 .566 29 Regression 9.463 .001
G_ 0.080X — 0.978 .885 58 Slopes 6.813 001 4.857*
A 0.086X — 1.218 .889 165

12584 (2, 99.1-128) Lama Stop-off; Eigenmann 1908 Exped. CAS IUM 12585 (2,
79-98) Rupununi Pan, opposite Massara Landing; Eigenmann 1908 Exped. CAS
IUM 12586 (2, 92.4-97.8) Essequibo R. at Rockstone; Eigenmann 1908 Exped.
CAS SU 21881 (1, 126) Lama Stop-off; Eigenmann 1908 Exped. USNM 66279 (1,
81.9) Rupununi Pan, opposite Massara Landing; Eigenmann 1908 Exped. ANSP
39799_-800 (2, 76.0-104) Rupununi R.; J. Ogilvie. CAS IUM 12579 (3, 66.9-107)
Konawaruk Pool; Eigenmann 1908 Exped. MCZ 30193 (1, 101) Essequibo R. at
Rockstone; Eigenmann 1908 Exped. ZMH 17821 (5, 64.8-120) Essequibo R. at

VOLUME 95, NUMBER 4 12>

Table 15.—Newman-Keuls Multiple Range Tests of significantly different slopes or intercepts for
regressions of morphometric data on body length in 3 populations of Potamorrhaphis guianensis.
Regressions and analysis of covariance from Table 14. (* = significant at 0.05).

P,—P, distance Intercepts Q for ranks

1. Guianas —3.19720 1-3 6.4825*
2. Amazon = 2 SI) 1-2 4.8271*
3. Orinoco —2.39392 2-3 1.9974
P.-C distance Intercepts Q for ranks

1. Orinoco 1.21725 1-3 Se QZ
2. Amazon 2.25079 1-2 0.4837
3. Guianas 3.90308 2-3 6.1374*
Interorbit Slopes Q for ranks

1. Orinoco 0.04490 1-3 4.8653*
2. Guianas 0.05559 1-2 Z Sks ey
3. Amazon 0.06284 2-3 3.6200*
Head width Slopes Q for ranks

1. Orinoco 0.04135 1-3 5.8550*
2. Guianas 0.05900 1-2 4.9729*
3. Amazon 0.06714 2-3 3.3266*
Pelvic fin length Slopes Q for ranks

1. Orinoco 0.05103 1-3 4.8574*
2. Guianas 0.08017 1-2 4.6335*
3. Amazon 0.08606 2-3 1.5534

Rockstone; 9 Nov 1938; W. Griem. NHMV uncat. (1, 68.2) Rupununi Pan op-
posite Massara Landing; Eigenmann 1908 Exped. NHMV uncat. (1, 40.4) Ru-
pununi R.; Haseman. NHMV uncat. (1, 94.3) Rupununi R.; 23 Feb 1913; Hase-
man. NHMV uncat. (6, 73.8-120) Rupununi R. BMNH 1972.7.27.1014-20 (6,
86.7-116) Rupununi R. at Karanambo; 2 Aug 1958; R. H. Lowe-McConnell.
BMNH 1972.7.27.1010—1013 (4, 56.8-72.3) Rupununi R., Pirara Stop-off; 1957;
R. H. Lowe-McConnell. MCZ 48560 (1, 124) Rupununi R., Agua Branc, tributary
of Manari R.; 8 May 1971; C. Hopkins. MCZ 48559 (2, 94.3-97.4) Rupununi R.,
Agua Branc, tributary of Manari R.; 20 Apr 1971; C. Hopkins. AMNH 15192 (10,
60.8—130) Essequibo R. at Rockstone; Mar 1938; Pinkus. DEMARARA RIVER,
GUYANA: 6 (68.6—133) from 3 collections. CAS IUM 12583 (2, 88.5-111) Wis-
mar; Eigenmann 1908 Exped. BMNH 1926.5.27.9-10 (2, 116-133) Demarara R.;
Imp. Bur. Ent. BMNH 1936.4.4.38-39 (2, 68.6-97.7) Wismar. NICKERIE RIV-
ER, SURINAM: 19 (80.2—136) from 5 collections. ZMA 105.814 (4, 80.2—118) 12
km WSW Standansie Falls; 4 Apr 1967; H. Nijssen—120. RMNH uncat. (N-1)
(1, 122) L tributary below Blanche Marie Falls; 16 Feb 1971; M. Boeseman.
RMNH (N-2) (2, 98.8-121) above Camp Standansie; 30 Jan 1971; M. Boeseman.
RMNH uncat. (N-3) (4, 86.0-136) Tjawassi Cr.; 7 Feb 1971; M. Boeseman. RMNH
uncat. (N-4) (8, 98.7-127) R tributary of Fallawatra, 35 mi from outlet; 2 Feb
1971; M. Boeseman. SARAMACCA RIVER, SURINAM: 4 (117-140) from 2
collections. ZMA 105.604 (2, 125—140) Toebaka Cr.; 2 Mar 1967; H. Nijssen—
110. ZMA 105.627 (2, 117—130) Kleine Saramacca R.; 27 Feb 1967; H. Nijssen—

736 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

106. SURINAME RIVER, SURINAM: 52 (89.0-—144) from 25 collections. ZMA
106.206 (1, 119) Marowijne Cr., 63 km S Afobaka; 22 Oct 1966; H. Nijssen—79.
ZMA 105.706 (1, 112) Suriname R., 1 km S Botopasi; 22 Mar 1967; H. Nijssen—
117. ZMA 105.785 (1, 93.5) Awala Cr., 1.5 km S Botopasi; 18 Mar 1967; H.
Nijssen—1I11. ZMA 105.703 (2, 101-112) Jenjee Cr., 7.5 km N Botopasi; 21 Mar
1967; H. Nijssen—114. ZMA 105.204 (5, 108-130) Sara Cr., Brokopondo, 27 km
S of village dam; 14 Oct 1966; H. Nijssen—76. ZMA 105.503 (11, 110-140) Mar-
chall Cr.; 8 Dec 1966; H. Nijssen—87. RMNH uncat. S-1 (1, 109) Kwambado
Kr., R. tributary of Sara Kr.; 22 Dec 1963; M. Boeseman. RMNH uncat. S-2 (2,
93 .8—99.3) Langatabbetje, upper Sara Kr.; 12-14 Dec 1965; G. F. Mees. RMNH
uncat. S-3 (1, 89.0) Langatabbetje, upper Sara Kr.; 11 Dec 1965; G. F. Mees.
RMNH uncat. S-4 (2, 112-120) Compagnie Kr.; 19 Dec 1965; G. F. Mees. RMNH
uncat. S-5 (3, 123-138) Jabokai Kr.; 18 Feb 1964. RMNH uncat. S-6 (1, 92.6)
Compagnie Kr.; 13 Apr 1965; G. F. Mees. RMNH uncat. S-7 (1, 125) Gran Kr.,
12 km from outlet; 21 July 1964; M. Boeseman. RMNH uncat. S-8 (1, 122) R.
tributary middle Gran Kr.; 30 July 1964; M. Boeseman. RMNH uncat. S-9 (3,
108—129) Amaniparicreek, R. tributary of Sara Kr.; 24 Feb 1964; M. Boeseman.
RMNH uncat. B-1 (2, 119-137) Brokopondo, Taproepa Kr.; 20 Jan 1964. RMNH
uncat. B-2 (2, 127-128) Brokopondo; 17 Dec 1963; M. Boeseman. RMNH uncat.
B-3 (1, 113) Brokopondo, Suriname R.; 12 May 1964; M. Boeseman. RMNH
uncat. B-4(1, 136) Brokopondo, Suriname R.; 2 May 1964; M. Boeseman. RMNH
2549 (1, 126) Brokopondo, Marchall Kr.; 28 Dec 1965; G. F. Mees. RMNH
uncat. B-6 (3, 120-144) Brokopondo, between Brokopondo and Afobaka; 13 Dec
1963; M. Boeseman. RMNH uncat. B-7 (3, 90.4—144) Brokopondo, Taproepa Kr.;
22 Dec 1963; M. Boeseman. RMNH uncat. B-8 (1, 96.3) Brokopondo; 10 Jan
1964; M. Boeseman. RMNH uncat. B-9 (1, 110) Brokopondo, Suriname R.; 13
Feb 1964; M. Boeseman. RMNH uncat. B-10 (1, 118) Brokopondo, Suriname R.;
5 Jan 1964; M. Boeseman. MARONI (=MAROWIJNE) RIVER, SURINAM—
FRENCH GUIANA: 7 (54.8—132) from 3 collections. ZMA 106.207 (2, 128-132)
Kamaloea Cr., 9 km SE outlet Gran Cr.; 24 Apr 1967; H. Nijssen—131. ZMA
108.334 (4, 54.8-82.1) Albina; 7 Aug 1960; H. Pijpers. USNM 211329 (1, 94.6)
Grand Santi; 6 Sept 1971; E. Remale. MANA RIVER, FRENCH GUIANA:
MNHN 833 (1, 123) La Mana; holotype of Belone scolopacina. AMAZON RIV-
ER: 264 specimens (33.7—180) from 83 collections. RIO SOLIMOES: 58 (33.7—
180) from 22 collections. CAS SU 50636 (3, 117—126) Colombia, Caqueta Prov.,
R. Orteguaza between Tres Esquinas and Solano; 12 Feb 1958. USNM 234943 (1,
119) Ecuador, Pastaza Prov., R. Bufeo, tributary of lower Borbonoza R.; Feb
1963; Olalla. USNM 163891 (2, 107-130) Ecuador, Pastaza Prov., Chichirota on
lower Borbonoza R.; R. Olalla. CAS IUM 15811 (1, 133) Peru, Loreto, Morona
R.; Oct 1920; W. R. Allen. CAS [UM 15815 (1, 96.8) Peru, Loreto, Pacaya R.;
Aug 1920; W. R. Allen. ANSP 73163 (2, 89.9-121) Peru, Loreto, Ucayali R. at
Contamana; 1937; W. C. Morrow. ANSP uncat. (5, 75.2—114) Peru, Loreto, R.
Nanay near Iquitos; 19 Oct 1955; Hohn. UMIM 1110 (1, 97.2) Peru, Loreto, near
Iquitos; 12 Sept 1954; L. R. Rivas. NHMV uncat. (2, 123-133) Peru, Loreto,
Iquitos; Steindachner. NHMV uncat. (4, 101-114) Peru, Loreto, Iquitos; Stein-
dachner. CAS SU 17272 (2, 102-136), 36849 (2, 107-114), 36850 (1, 120), 36851
(1, 90.8), 36852 (5, 88.0-120), 36853 (1, 91.0), 58700 (1, 33.7), Peru, Loreto, vi-
cinity of Pebas in Rio Ampiyaca, Tuye Cano, etc.; 1936-41; W. G. Scherer. ZMH

VOLUME 95, NUMBER 4 137)

17711 (9, 109-143) Peru, Loreto, between Iquitos and Leticia, 1937; H. Pietsch.
ZMH 17673 (1, 110) Peru, Loreto between Iquitos and Leticia; 1936; H. Pietsch.
UAIC 4165 (2, 96.2—103) Colombia, Amazonas, tributary to Loreto Yacu R.; 17
June 1971. MCZ 2760 (3, 68.7—106) Brazil, Tabatinga (=Sapurara); Thayer Exped.
MCZ 8799 (8, 85.0-147) Brazil, R. Hyavary (=Javari) near Tabatinga; Thayer
Exped. MIDDLE AMAZON, BRAZIL: 10 (103-180) from 5 collections. MCZ
2758 (3, 103-180) Brazil, Jutai; Thayer Exped. SMF 6714 (2, 104-125) Brazil, R.
Solimoes, Iguarapé Preto; 14 Mar 1961; H. Schultz. USNM 222298 (3, 132-172)
Brazil, R. Tefé, Juruparu, half way up river; Aug 1979; M. Goulding. MCZ 8800
(1, 118) Codajas; Thayer Exped. MCZ 52601 (1, 118) Brazil, Amazonas, Rio
Solimoes, Ilha de Morta opposite Parana de Janauaca; 3°25’S, 60°21'W; 20 Nov
1976; W. L. Fink 76-1. RIO NEGRO, BRAZIL: 42 (65.2—134) from 6 collections.
MCZ 52600 (14, 75.3—134) Brazil, tributary of Rio Cuieiras 3 km from Rio Negro;
2°50’S, 60°35’W; 27 Nov 1976; W. L. Fink 76-5. NHMV uncat. (1, 121) Brazil,
R. Branco at Boa Vista; 1912; Haseman. NHMV uncat. (1, 105) Brazil, R. Ca-
vame, tributary of R. Branco at Boa Vista. IRSNB uncat. (1, 110) Brazil, Ar-
quipélago das Anavilhanas, sta. 179; 18 Nov 1967. IRSNB uncat. (1, 105) Brazil,
Arquipélago das Anavilhanas, sta. 180; 19 Nov 1967. USNM 228955 (24, 65.2-
115) Guyana, Pirara R. area; 31 Jan 1953. LOWER AMAZON, BRAZIL: 105
(56.0—-165) from 39 collections. MCZ 2754 (5, 110-259) Manacapuru; Thayer Exped.
ZMH 16032 (1, 134) Manacapuru; 1924; W. Ehrhardt. ZMH 17483 (1, 118) Ma-
_naus. BMNH 1893.4.24.40 (1, 124) Manaus; J. C. Antony. FMNH 94710 (2, 101-
135) Manaus; C3156; 29 Nov 1909. BMNH uncat. (1, 125) Lago Doro near Ma-
naus; Nov 1972. CAS SU 52682 (5, 85.1—134) Manaus, Igarapé de Mai Grais; 25
Dec 1924; C. Ternetz. USNM 179527, 179528, 179529 (14, 67.3—132) Rio Urubu
25 mi from Itacoatiara; H. Axelrod. CAS SU 52685 (2, 122-124) Igarapé da Mae
Joana; C. Ternetz. CAS uncat. (1, 99.2) Igarapé da Mae Joana; 25 Dec 1924; C.
Ternetz. 3 CAS uncat. (5, 93.8-137) Lago Grande into Amazon; Oct-Nov 1924;
C. Ternetz. IRSNB uncat. (1, 112) Igarapé Matapi, Lago Matapi, Trombetas R.,
sta. 136; 2 Dec 1964. MCZ 8770 (1, 136) Villa Bella above Obidos, L. Agassiz;
Thayer Exped. MCZ 8798 (2, 92.5—92.8) Villa Bella above Obidos, L. Agassiz;
Thayer Exped. MCZ 1036 (1, 122) Obidos; Jeffreys. MCZ 8801 (1, 109) Obidos,
Thayer Exped. MCZ 2757 (7, 56.0-122) Obidos, Thayer Exped. IRSNB uncat.
(3, 134-165) Bras du R. Cururu, mission franciscaire du Cururu, Haut Tapajox,
sta. 129; 21 Nov 1964. CAS SU 1926 (2, 131) Itaituba; Agassiz. CAS IUM 6959
(3, 94.0-106) Itaituba Bay; C. F. Hartt. MZUSP 8512 (5, 74.0—93.0) Santarém,
tributary of R. Mapiri; 25 Dec 1963. CAS SU 64332 (1, 103) and 3 CAS uncat.
(7, 108-137), Santarém market; Sept-Oct 1924; C. Ternetz. NHMV uncat. (4,
95.5—123) Santarem; Haseman. MCZ 1028 (1, 110) Santarém; J. C. Fletcher. MCZ
8787 (1, 129) Santarém; Thayer Exped. MCZ 8806 (1, 118) Santarém; 1873; C.
Linden. FMNH 94711 (1, 88.0) Santarém; C3284; 15 Dec 1909. MCZ 4682 (2,
133) Porto do Moz; Thayer Exped. MCZ 8788 (18, 114-157) Gurupa; Agassiz.
NHMYV 3 uncat. lots (5, 140-163) Gurupa; Steindachner. S. AMAZON TRIBU-
TARIES, BRAZIL: 14 (65.9-119) from 3 collections. USNM 201436 (10, 72.4—
118) upper Rio Xingu, Mato Grosso; H. Schultz. USNM 191595 (2, 112-119) R.
Araguaia near Aruana; 1960; H. Axelrod. CAS IUM uncat. (2, 65.9-86.1) Tocan-
tins R.; 26 Apr 1924; C. Ternetz. PARA REGION, BRAZIL: 38 (82.9-160) from
10 collections. BMNH 1849.11.9.59-61 (3, 131-154) Capim R.; syntypes Belone

738 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

taeniata. MCZ 46067 (3, 147-152) R. Apeu, Boa Vista, Castanhal, Para; July
1965; N. Menezes. MCZ 46065 (8, 116-157) Igarapé Icatu, Belem; July 1965; N.
Menezes. MCZ 46066 (1, 107) R. Arari, Ilha da Marajo; July 1965; N. Menezes.
MCZ 46075 (2, 82.9-124) Igarapé Paracuri, Icoaraci; July 1965; N. Menezes.
NHMV uncat. (1, 116) Para; Mus. Goldi; Steindachner. NHMV uncat. (1, 146)
Lema bei Para; 1913; Haseman. NHMV uncat. (3, 105-151) Sema, Para; 6 Aug
1913; Haseman. MZUSP uncat. (13, 113-160) Para, Belém, Utinga. MZUSP un-
cat. (3, 151-159) Igarapé Apeu, Boa Vista; July 1965; N. Menezes.

Potamorrhaphis eigenmanni A. de Miranda Ribeiro
Fig. 4d

Belone taeniata (not of Gunther, 1866) Perugia, 1891:654 (Villa Maria, Mato Gros-
so).—Boulenger, 1896:37 (Descalvados, Mato Grosso).

Belone (Potamorrhaphis) taeniata (not of Gunther, 1866) Perugia, 1897:26 (Rio
Madidi, tributary of Rio Madeira).

Potamorrhaphis guianensis (not of Schomburgk, 1843) Eigenmann, McAtee, and
Ward, 1907:143 (Rio Paraguay at Tuyuyu).—Pearson, 1924:52 (Lake Rogoagua
and near Reyes, Rio Beni basin, Bolivia), 58 (SO0—1500 ft. altitude).—Pearson,
1937:112 (present in both the Beni-Mamoreé and Paraguay basins).—Fowler,
1940b: 102 (listed from the Madeira-Mamoré and Paraguay water sheds after
Perugia 1897, and Pearson 1924).—Pozzi, 1945:264, 276 (Rio Paraguay).—
Luengo, 1972:22 (Mato Grosso).

Potamorrhaphis eigenmanni A. de Miranda Ribeiro, 1915:I-13 (original descrip-
tion; Porto Esperidiao, Rio Jauru, Mato Grosso).—P. de Miranda Ribeiro, 1953:
395 and 1961:7 (reference to original description and types).—MacDonagh,
1938: 188-190 (comparison of types of B. taeniata and specimens from Manaus
and the Demarara River with 7 specimens from the Paraguay River by Dr.
Trewavas, BMNH; B. taeniata D 32-34, A 28-31 vs. P. eigenmanni D 28-31,
A 24-27). Pozzi, 1945:264, 276 (Rio Parana and R. Paraguay).—Collette, 1974a:
34 (listed as nominal species).—Castello et al., 1978:134 (description; middle
Parana River, Argentina; fig. 8).

Potamorrhaphis taeniata (not of Gunther, 1866) Pozzi, 1945:264, 276 (in part,
Rio Paraguay).

Diagnosis.—Most similar to P. guianensis from which it differs most markedly
in lower numbers of dorsal- and anal-fin rays, vertebrae, and predorsal scales
(Tables I-7).

Types.—Potamorrhaphis eigenmanni A. de Miranda Ribeiro, 1915. Lectotype:
MNRJ 1343A (95.0 mm); Brazil, Mato Grosso, Rio Jauru at Porto Esperidiao; A.
de Miranda Ribeiro; selected by P. Miranda Ribeiro (1953:395). D 29; A 26; P, 8
(?); P,—-P. distance 47.5 mm; P.—C distance 45.3 mm. Paralectotype: MNRJ 1343
(95.0 mm).

Geographic variation.—There are two major populations of P. eigenmanni,
located in the Paraguay-Parana drainage basin and in the upper Rio Madeira, a
tributary of the Amazon. Material is available from three tributaries of the Ma-
deira, the Mamoré, Guaporé, and Beni rivers (Fig. 5). One specimen of P. ei-
genmanni comes from further downstream in the Madeira, below the junction

VOLUME 95, NUMBER 4 739

: 80° ae | 50° 40°
3S eal
os ag ag {PILE [SB,QRINOCO 102
ais a GUIANAS

< |
as x éw Branco NG SMAMAZON

(0) S¢ Zo (0)
A, Negro Seo. AeA |
SPQ ei i \ \(?
Solimées NS og ie 4
‘Madeira * Tapajos
i 2 / \ ) |
N. Ucayali
1 (Xingu
10 eS Araguaia— >
a By
OD OF 9d A
20 i ; =
a Parana
e P. guianensis es LS
© P. petersi ~ ;
P_ eigenmanni :
Vy
30° j

LA PLATA | :
50 40°

Fig. 5. Distribution of the 3 species of Potamorrhaphis based on specimens examined.

with these three tributaries. ANOVA comparisons of seven meristic characters
were made among the five populations of P. eigenmanni (Table 16). The four
Madeira populations were then combined and this combined population was tested
against the Paraguay population (last 2 lines in Table 16).

In comparing the five populations, statistically significant differences between
means were found for five of seven characters, all tested except numbers of
pectoral-fin rays and predorsal scales (Table 16). There is a general pattern of
decreasing counts from the Madeira (one specimen) to the Mamoreé, Guaporé,
Beni and finally the Paraguay. In some cases, population means do not differ
from each other: Madeira and Mamoré (anal-fin rays); Mamoré, Guaporé, and
Beni, (precaudal vertebrae); Guaporé and Beni (total vertebrae); Guaporé, Beni,
and Paraguay (caudal vertebrae); Beni and Paraguay (dorsal- and anal-fin rays).
Combining all the Madeira populations and testing their mean against that of the
Paraguay population showed statistically significant differences in all seven char-

740 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 16.—Comparison of means for meristic characters for 5 populations of Potamorrhaphis
eigenmanni (* indicates mean is significantly different from adjacent mean by SNK, alpha = 0.05).

Fin rays Vertebrae Scales

Population n Dorsal Anal Pectoral Precaudal Caudal Total Peden
Madeira 02> 933% 280 lara (8 ae = = 101.0
Mamoré GE10) Silks ies 7.8 STW, By ao) 70.0* yO) S)
Guaporé 30-33 = 30.2* 26.4* 7.0 Bie ae Sey 68.6 |, 97.4
Beni NS) ee 857) || 25 as Ud 3 22 30.6 ¢* 67.9 93.4
Paraguay 49-66 29.0 25.4 Uc”? 36.4* 30.3 66.7* 90.6

Summary
Total Madeira 50259) 30107 ZOO VO Me aiea Sess 68.6* SY2*

Paraguay 49-66 29.0% 25.4* D2 SOr4E 307 66.7* 90.6*

acters (Table 16). The mean for each character for the Paraguay population is
less than that for the Madeira population.

Morphometric data for the four Madeira populations were combined and tested
against the Paraguay populations by ANCOVA. Statistically significant differ-
ences were found in seven of the 12 morphometric characters (Table 17). Slopes
were different for P,—-P, distance, postorbital distance, and preopercle length.
Intercepts were different for orbit length, interorbital width, head depth, and head
width.

Material examined.—184 specimens (31.5—132 mm BL) from 46 collections.
UPPER MADEIRA RIVER, BOLIVIA AND BRAZIL: 91 (45.0—132) from 17
collections. CAS IUM 17254 (2, 78.5—85.0) Bolivia, R. Beni near Reyes; Oct 1921;
N. E. Pearson. USNM 86775 (2, 104-111) Bolivia, R. Beni, Lake Rogoagua; Nov
1921; N. E. Pearson. CAS 14376 (5, 72.2—126) and FMNH 88483 (1, 111) Bolivia,
Lake Rogoagua; Nov 1921; N. E. Pearson. UMMZ 66427 (5, 45.0—-117) Bolivia,
Lake Rogoagua. MSNG 13962 (1, 132) Bolivia, R. Madidi, tributary of R. Ma-
deira; 1893; L. Balzan. AMNH 39823 (55, 62.5—125) Bolivia, Dept. Beni, R. Itenez
(Guaporeé), 10 km SE Costa Marques, Brazil; 10 Sept 1964; R. M. Bailey and R.
Ramos. SMF 11866 (1, 89.1) Bolivia, Mamoré; C. A. Hahn. NHMV 11125 (2,
80.0-82.5) and NHMV 1126 (2, 88.4—90.4) Bolivia, R. Guaporé; C. Natterer.
FMNH 94702 (2, 89.7—94.2) Bolivia, San Joaquin on R. Machupo; C2935; 4 Sept
1909. FMNH 94703 (1, 97.4) Bolivia, Maciel, R. Guaporé; C2830; 23 July 1909.
FMNH 94705 (1, 70.0) Bolivia, Maciel, R. Guaporé; C2845; 28 July 1909. FMNH
94704 (1, 109) Bolivia, Maciel, R. Guaporé; C2881; 3 Aug 1909. FMNH 94707 (1,
51.5) Brazil, Bastos, R. Alegre into R. Guaporé; 8 mi S Villa de Mato Grosso;
C2795; 26 June 1909. IRSNB 19588 (6, 86.2—127) Brazil, Rondonia, Guajara-
Mirim, Sta. 184; 26 Nov 1967. IRSNB 19589 (3, 108-112) Brazil, Rondonia, Gua-
jara-Mirim, Sta. 185; 26 Nov 1967. IRSNB uncat. (1, 136) Brazil, Rond6nia, Porto
Velho, Igarape dos Milagres; Sta. 181; 24 Nov. 1967. PARAGUAY RIVER: 91
(31.5—123) from 27 collections. MNRJ 1343 (2, 95.0) Brazil, Mato Gross, R. Jauru,
Porto Esperidiao; A. de Miranda Ribeiro, lectotype and paralectotype of Pota-
morrhaphis eigenmanni. MNRJ 1344 (1, 106) Brazil, Mato Grosso, Lagoa de
Caceres, Corumba; 1913; F. L. Hoehne. FMNH 94708 (1, 97.8) Campos Alegre,
R. Jauru, C2759. FMNH 94709 (2, 95.5—97.8) Sao Luiz de Caceres; C2758; 27

VOLUME 95, NUMBER 4 741

Table 17.—Analysis of covariance of regressions of morphometric data against body length for
populations of Potamorrhaphis eigenmanni from the Madeira (M) and Paraguay (P) drainages.

Character Pop. Y= CD N F P
Head length M 0.598X — 1.904 .954 40 Regression 0.203 .654
2 0.617X — 3.768 959 32
Snout length M 0.449X — 3.309 945 40 Regression 1.344 250
P 0.456X — 4.667 929 32
P,—P, M 0.480X — 1.212 .976 48 Regression 7.291 .008
P 0.521X — 2.024 .988 55) Slopes 9.297 .003
P.-C M 0.486X — 0.127 .980 48 Regression 3.510 .064
P 0.456X + 2.273 .984 55
Postorbit M 0.101X + 1.452 .905 48 Regression 12.496 .001
P 0.117X + 0.425 .942 Sil Slopes 6.836 .010
Preopercle M 0.058X + 0.533 .862 48 Regression 9.423 .003
P 0.066X + 0.057 .947 57 Slopes 4.200 043
Orbit M 0.036X + 0.944 915 48 Regression 11.302 .001
P 0.041X + 0.796 .847 56 Slopes 1.911 .170
Intercepts 20.305 .001
Interorbit M 0.582X — 0.540 .949 48 Regression 26.476 .001
P 0.063X — 0.572 931 56 Slopes 2.053 155
Intercepts 49.875 0
Head D M 0.070X — 0.627 .962 48 Regression 69.469 0
P 0.071X — 0.220 .970 56 Slopes 0.252 .617
Intercepts IB Ses57) 0
Head W M 0.056X + 0.129 57 48 Regression 16.623 O01
P 0.055X + 0.571 .932 56 Slopes 0.425 516
Intercepts 32.683 .001
J M 0.115X + 0.078 839 46 Regression 0.851 359
P 0.112X + 0.550 919 48
| Eee) M VOWS = Ueat) .876 48 Regression 0.049 .825
P 0.078X — 0.679 .901 55

May 1909. FMNH 94706 (14, 36.0-107) Paraguay, Puerto Suarez; C2690; 6 May
1909. MNM 641 (23, 57.0-89.4) Brazil, Mato Grosso, Laguna Gaibu near Amolar;
25 Oct 1955. MSNG 39849 (1, 78.8) Brazil, Mato Grosso, Villa Maria; L. Balzan.
FMNH 70389 (1, 64.5) Brazil, Mato Grosso, Descalvados; July 1926; Schmidt
and Sanborn. FMNH 70390 (4, 76.7—104) Brazil, Mato Grosso, Descalvados; July
1926; Schmidt and Sanborn. FMNH 70391 (6, 66.5—105) Brazil, Mato Grosso,
Conceicao; 24 July 1926; Schmidt and Sanborn. FMNH 70392 (1, 89.1) Brazil,
Mato Grosso, Conceicao; 10 Aug 1926; K. P. Schmidt. CAS 10267 (1, 90.2) Brazil,
Mato Grosso. MACN uncat. (2, 86.0-88.0) Brazil, Mato Grosso; Burmeister
Exped. BMNH 1895.5.17.259-60 (2, 72.3-81.3) Brazil, Mato Grosso; C. Ternetz.
BMNH 1900.4.14.87-90 (4, 68.6—-92.8) Carandasinho; A. Boretti. MSNG 14021
(2, 64.0-98.4) Paraguay, Puerto 14 Mayo; 1900; F. Silvestri. NHMYV uncat. (2,
102-123) Buchten Paraguay; Natterer. NHMV uncat. (2, 96.0—105) Buchten Par-
aguay; Natterer. NHMV 11127 (2, 82.7-89.8) Paraguay; Natterer. NHMV 11128
(4, 68.8-83.3) Paraguay; Natterer. BMNH 1910.5.26.51 (1, 126) Paraguay; Tudor.

742 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 18.—Analysis of covariance of regressions of morphometric data on body length for the 3
species of Potamorrhaphis: P. petersi (P), P. guianensis (G), and P. eigenmanni (E). (* = significant

at 0.05).

Character Pop. Y= CD N P Q
Head length Pa OFS 45 Xe 2962 aeoral 10 Regression 12.985 <.001
G 0S80xe = 063 2910" 1585 Slopes 1.010 366
1 OLCOK = 2p S57 72 Intercepts 24.741 0 8.832*
Snout length P 0.410X — 3.924 .962 10 Regression 9.878 <.001
Gr 0142306507443 2890" S385) Slopes 1.184 308
E 0.454K — 4.166 .937 (2 intencepus 18.382 0 1.970
P,-P, | en Sh), Ge Ey, 0) 16 Regression 63.152 0
Ge 0PSIOXe 278225 982 SS S1OpES 4.150 .016 32s
E 0.499X — 0.371 .981 103
P,—C PO SOX we Sool 16 Regression 70.964 0
Ge UAC ae 2S BK Sleysres 5.868 .003 4.489*
I) OLDS MO | SS2 G8
Postorbit P 0.097X + 0.417 .940 16 Regression 33.426 0
G O:110X = 05568) 2931 266 Slopes 1.786 .169
1 UNO se Qs) 105 Intercepts 64.460 0 14.047*
Preopercle 1 OSI. = O30 9 Osx 16 Regression SGIS 0
G 0.060X + 0.167 .910 260 Slopes 0.437 .646
E 0.061X + 0.403 .900 105 Intercepts 64.758 0 SES 95
Orbit I Bay sb ORI Wile! 16 Regression 21.877
G 0.038X + 1.060 .873 267 Slopes 0.548 578
Ee) 02037 X60: 995 e843 104 += Intercepts 43.081 11.407*
Interorbit IP YOS52% S358 {Sls 16 | Regression 54.788 0
Ge? 0063x—— 15025 2925 3267 Se Slopes BE322 .037 2.500
E 0.058X — 0.330 .905 104
Head D I OWES SO AS 16 Regression 39.884 0
G 0.070X — 0.610 .935 247 Slopes 1.398 248
E 0.067X — 0.071 .920 104 Intercepts 77.768 0 15.524*
Head W 2 OOD CS TS S07 16 Regression 12.956 <.001
G 0.066X — 0.792 .908 247 Slopes 12.862 <.001 6.453*
i) OM SCaeO,s5D > LLzZ 104
Pale P Ont Xe 1074602933 14 Regression 13.834 <.001
G 0.126X — 0.734 .886 223 #£Slopes 3.382 .035 1.743
J OSI ee O'S cei) 94
Rosle P 0.086X — 2.237 .967 16 Regression 8.906 .0002
G 0.087X — 1.441 .886 252 Slopes 2.990 052
Ja, OOD = OOAS — ts327/ 103 Intercepts 14.583 <.001 Spo lo*

UMMZ 207843 (1, 102) Paraguay, R. Aquidaban at Paso Horqueta ca. 24 km
NNW of Loreto; 5-6 Sept. 1979; J. Taylor et al. UMMZ 205601 (1, 48.1) Para-
guay, overflowing inlet along E shore of R. Paraguay ca. 1 km S from Puente
Remanso bridge; 21 May 1979; J. Taylor et al. UMMZ 207252 (1, 55.8) Paraguay,
R. Montelindo ca. 155.8 km NW of Benjamin Aceval; 16 Aug 1979; G. R. Smith
and J. N. Taylor. UMMZ 205885 (1, 31.5) Paraguay, E shore of R. Paraguay; 11
June 1979; R. M. Bailey et al. UMMZ 207901 (4, 54.0-123) Paraguay, R. Aqui-
daban at Paso Horqueta ca. 24 km NNW of Loreto; 6 Sept 1979; J. Taylor et

VOLUME 95, NUMBER 4 743

al. UMMZ 206673 (5, 68.4-102) Paraguay, R. Aquaray Guazei 5.4 km S of junc-
tion with road W to San Pedro; 22 July 1979; J. Taylor. PARANA RIVER,
ARGENTINA: MACN 4291 (1, 77.5) R. Parana; 12 Nov 1895.

Discussion

As hypothesized previously (Collette 1966), Potamorrhaphis probably evolved
from inshore marine needlefishes similar to Strongylura, having three pairs of
upper pharyngeal tooth plates (UP,, UP;,,, and UP;), a truncate caudal fin, and
unforked pelvic fins smaller than the pectoral fins. Potamorrhaphis and Belonion
share such specialized characters as a rounded caudal fin, few large teeth on the
pharyngeal bones, pectoral-fin rays reduced to eight or fewer, nasal barbel elon-
gate, and an association of the expanded first neural spine with the supraoccipital
crests and exoccipital flange. Potamorrhaphis diverged from Belonion by elon-
gating the posterior part of its body and increasing the numbers of caudal ver-
tebrae and dorsal- and anal-fin rays. The evolutionary line leading to Belonion
continued to specialize by reduction of body size, number of vertebrae, bran-
chiostegal rays, and pectoral-fin rays and by the loss of a pharyngeal tooth plate
(UP3).

The three species of Potamorrhaphis form a series with increasing numbers of
dorsal- and anal-fin rays, vertebrae (precaudal, caudal, and total), and predorsal
scales from P. eigenmanni to P. guianensis to P. petersi. The differences in these
‘characters are greater between P. petersi and P. guianensis than between the
latter and P. eigenmanni. In comparison with presumably less specialized needle-
fishes, such as Strongylura, P. eigenmanni appears to be the least specialized of
the three species, P. petersi the most. Geographically, the three species form a
similar series from south to north with P. eigenmanni concentrated in the Para-
guay-Parana system, P. guianensis distributed throughout the Amazon and the
Guianas, and P. petersi concentrated in the Upper Rio Orinoco. Apparently, P.
eigenmanni has invaded the Amazon through the Mato Grosso into the Upper
Rio Madeira in Bolivia and Brazil. Similarly, P. petersi seems to have moved
through the Rio Casiquiare into the upper Rio Negro in Brazil. I know of no
sympatric occurrences of P. eigenmanni with P. guianensis, although the latter
does occur farther downstream in the Rio Madeira (Fig. 5). Similarly, P. petersi
occurs in the upper Rio Negro and P. guianensis farther downstream. Some
intergradation may have taken place between P. eigenmanni and P. guianensis
in the Rio Madeira and between P. petersi and P. guianensis in the Rio Negro
as suggested by the frequency distributions of meristic characters (Tables 1-2,
4—7).

This south-to-north pattern, from the Parana to the Amazon to the Orinoco, Is
similar to the recent interpretation of the origins of the parasitic helminths of
South American freshwater stingrays (Potamotrygonidae) by Brooks, Thorson,
and Mayes (1981). Both Potamorrhaphis and the Potamotrygonidae are absent
from the Rio Sao Francisco system of eastern Brazil. However, the Potamotry-
gonidae range west into the Maracaibo and Magdalena drainages from which
Potamorrhaphis is absent. Brooks et al. further hypothesize that the relationships
of the Potamotrygonidae are not with Atlantic dasyatids but rather with Pacific
Urolophus. While I have no morphological information that would support the
origin of Potamorrhaphis from Atlantic or Pacific Strongylura, | should note that

744 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

a parasitic ergasilid copepod, Ergasilus orientalis, was reported from two species
of needlefishes, Potamorrhaphis guianensis from the Amazon and Strongylura
incisa from northern Australia (Cressey and Collette 1970) as well as from a goby
and an atherinid from Japan. If these copepod populations really are conspecific,
this distribution would tend to support the hypothesis of Brooks et al. (1981).

Recognition of P. eigenmanni as a distinct species confirms the total number
of species in the family as 32 and the known neotropical species as nine in four
genera: Potamorrhaphis (3), Belonion (2, see Collette 1966), Pseudotylosurus (2,
see Collette 1974b), and Strongylura (2, S. hubbsi in the Usumacinta Province
of México and Guatemala, see Collette 1974c, and S. fluviatilis west of the Andes
in Colombia and Ecuador). I have previously (Collette 1974c:618) called attention
to the large number of freshwater needlefishes compared to halfbeaks in the
neotropics and the reverse situation in the freshwaters of the Indo-Australian
region. The nine neotropical freshwater species of belonids comprise 28.1% of
the species in the family; the two Indo-Australian freshwater species 6.3%. The
two neotropical freshwater halfbeaks, Hyporhamphus mexicanus Alvarez in the
Usumacinta Province of Middle America and H. brederi (Fernandez Yépez) in
the Orinoco and Amazon basins, comprise 2.9% of the approximately 70 species
in the family; the approximately 14 Indo-Australian freshwater species (in the
genera Dermogenys, Nomorhamphus, Hemirhamphodon, Hyporhamphus, and
Zenarchopterus) 20.0%. Historical, evolutionary, or ecological explanation of this
complementarity in distribution is needed.

Acknowledgments

I gratefully acknowledge the assistance and cooperation of the curators whose
institutions house material of Potamorrhaphis: Reeve M. Bailey (UMMZ); Marie-
Louise Bauchot (MNHN); M. Boeseman (RMNH); Herbert T. Boschung (UAIC);
James E. Bohlke (ANSP); Antenor Leitao Carvalho (MNRJ); Plutarco Cala (IN-
DERENA); William N. Eschmeyer (CAS); William L. Fink (MCZ); J. P. Gosse
(IRSNB); Robert K. Johnson (FMNH); P. Kahsbauer (NHMYV); M. Klappenbach
(MNM); W. Klausewitz (SMF); W. Ladiges (ZMH); Rogelio B. Lopez (MACN);
Francisco Mago Leccia (MBUCV); Naercio M. Menezes (MZUSP); Han Nissen
(ZMA); Donn E. Rosen (AMNH); Enrico Tortonese (MSNB); Haroldo Travassos
(MNRJ); and P. J. P. Whitehead (BMNH). M. Goulding provided fresh material
of P. petersi and C. E. Dawson facilitated loan of MZUSP specimens. Figure |
was prepared by Ruth Gibbons. Figures 2, 3, and 4c were drawn by Mildred H.
Carrington and Figures 4a, b, d by Keiko Hiratsuka Moore. Radiographs were
taken by George Clipper and Ruth Gibbons. Data were processed by Joseph L.
Russo and Ruth Gibbons. Drafts of the manuscript were reviewed by Naercio A.
Menezes, Joseph L. Russo, William L. Fink, Richard P. Vari, and Austin B.
Williams.

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PROC. BIOL. SOC. WASH.
95(4), 1982, pp. 748-762

A REVISION OF THE GENUS SIPHONOSOMA
(SIPUNCULA)

Edward B. Cutler and Norma J. Cutler

Abstract.—The sipunculan genus Siphonosoma (Spengel, 1912) is critically re-
viewed as the result of an examination of all existing type-specimens. The ratio-
nale for the separation into three subgenera is considered and found to be insup-
portable; the subgenera are discarded. Each of the 22 nominate species is
considered: ten remain as valid species, nine are reduced to junior synonyms,
two are considered incertae sedis and one species inquirendum. The genus is
redefined, the geographic distribution of the ten species is given and a key to
these species is presented.

Fisher (1950:805) divided the genus Siphonosoma into three subgenera which
he said were “‘fairly natural.’ It is not clear how he determined this naturalness
but he used the presence/absence of transverse dissepiments and numerous rectal
caecae to create his three groups. He did not give any rationale for selecting these
characters in this or subsequent papers.

The spelling of one subgenus, Dasmosiphon, was incorrectly changed to Da-
mosiphon in Stephen and Edmonds (1972), and that incorrect spelling has been
used by a few authors. In Stephen and Edmonds (1972) S. hataii was submerged
under §. carolinense, S. billitonense under S. edule, and S. formosum under S.
cumanense. The three species remaining in this subgenus were given considerable
attention and in Cutler and Cutler (1979) S. carolinense was submerged under S.
cumanense. Cutler, Cutler, and Nishikawa (in press) propose that S. edule and
S. cumanense are conspecific. Cutler (in press) petitioned the International Com-
mission on Zoological Nomenclature that, since S. edule and S. cumanense are
conspecific, the junior name, S$. cumanense, be conserved because it is the most
widely used and better known. Therefore, this subgenus now has one species, S.
cumanense.

The subgenus Hesperosiphon had three species assigned to it by Fisher (1950).
A new species, S. marchadi Stephen, 1960, was added and S. crassum has been
synonymized with S$. vastum (Stephen and Edmonds, 1972). The similarities be-
tween §. vastum and S. parvum are striking as noted by Stephen and Edmonds
(1972:57) but they retained three species. The subgenus Siphonosoma (sensu
stricto) has 16 species assigned to it in Stephen and Edmonds (1972, five of these
known only from a holotype).

Of the two characters selected by Fisher (1950) when he erected the subgenera,
one (transverse dissepiments) has been analysed in a large localized population
of S. cumanense from the Malagasy Republic. It was concluded that these thin,
fragile sheets of tissue are ephemeral and inconsistently present, ‘‘this is probably
the most variable of the variables in these worms” (Cutler and Cutler 1979:948).
It should not be used for species characterization, much less for a subgenus. The
second character, many rectal caecae, is not a ‘“‘natural’’ character if that implies
wide occurrence in the phylum; it is rare and probably of minor phylogenetic

VOLUME 95, NUMBER 4

Table 1.—Proposed status of species in this article.

Former status

Subgenus Dasmosiphon

Siphonosoma carolinense
Siphonosoma cumanense
Siphonosoma edule

Subgenus Hesperosiphon

Siphonosoma marchadi
Siphonosoma parvum
Siphonosoma vastum

Subgenus Siphonosoma

Siphonosoma amamiense
Siphonosoma arcassonense
Siphonosoma australe
Siphonosoma boholense
Siphonosoma bonhourei
Siphonosoma dayi
Siphonosoma eniwetoki
Siphonosoma funafuti
Siphonosoma ingens

Proposed status

Siphonosoma cumanense
Siphonosoma cumanense
Siphonosoma cumanense

Siphonosoma cumanense
Siphonosoma vastum
Siphonosoma vastum

Siphonosoma funafuti
Siphonosoma arcassonense
Siphonosoma australe
Siphonosoma boholense
incertae sedis
Siphonosoma dayi
Siphonosoma rotumanum
Siphonosoma funafuti
Siphonosoma ingens

Siphonosoma joubini incertae sedis

Siphonosoma mourense Siphonosoma mourense
Siphonosoma novaepommeraniae Siphonosoma cumanense
Siphonosoma pellucidum species inquirendum
Siphonosoma pescadolense Siphonosoma australe takatsukii
Siphonosoma rotumanum Siphonosoma rotumanum
Siphonosoma takatsukii Siphonosoma australe takatsukii

significance as judged by outgroup comparison. As a substitute we considered
using a primitive character occurring in other families i.e., contractile vessel villi,
as an indicator of related species within this genus, but are not proposing any
action at this time. However, we do propose returning to the concept of a genus
with no subgenera, now containing ten species.

In this paper we first comment on all the species considered valid in Stephen
and Edmonds (1972) in the order used therein. The literature references in this
part include only original findings (not review or repeat references). We will then
redefine the genus and present a key to the retained species. Our conclusions are
summarized in Table 1.

Two troublesome morphological characters.—TYhe term ““hook’’ has been ap-
plied to a variety of epidermal structures in this phylum. Within this genus there
exists an unusual range of structures. At one end of the continuum we find five
species with nothing but secretory papillae on the introvert (S. boholense, S.
cumanense, S. funafuti, S. ingens, and S. mourense). At the other end are two
species with typical, standard, large, pointed hooks (S. australe and S. vastum)
(Fig. 1C). Siphonosoma dayi and S. arcassonense have rings of chitinized tu-
bular, scalelike papillae which lie flat along the introvert but are arranged like,
and are probably homologous to, hooks (Fig. 1A & B). Siphonosoma rotumanum
has short, not sharply pointed hooks closely associated with a large papillum
(Fig. 1D).

750 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. !A-H. Hooks and contractile vessel of several Siphonosoma species to show variations
present within this genus (see text for discussion): A, S$. dayi, scale-like hooks viewed from above;
B, S. dayi, hooks, side view, length ranges from 0.13—0.17 mm; C, S. australe, hooks, height 0.19-
0.21 mm; D, S. rotumanum, hooks and associated papillae, height of hooks 0.1—-0.2 mm;.E, Ventral
retractor muscles (VR), digestive system and contractile vessel (CV) to show position of latter vessel
(V—ventral nerve cord; R—rectum; E—esophagus); F, Dorsal view of contractile vessel as it some-
times appears in contracted state with accordian pleats or folds; G, Side view of contractile vessel
as it Sometimes appears with bulbous swellings, bumps, or vesicles; H, Side view of contractile vessel
with true villi as in S$. cumanense. 1, Siphonosoma ingens, internal posterior end of trunk showing
the tubular, fusiform glandular bodies which are also present in S$. arcassonanse, (S—spindle muscle;
V—ventral nerve cord.)

Another character which requires careful examination and a clear understand-
ing of terms is the contractile vessel villi (Fig. 1E). The tubular contractile vessel
may be folded like a compressed accordion (Fig. 1F) or have bulbous pouches
or vesicles (Fig. 1G) along some portion of it but these are not villi. Villi are
generally digitiform or clavate with a small diameter, and the length clearly ex-
ceeds the width. They are outgrowths from, not swellings of, the underlying
vessel (Fig. IH).

VOLUME 95, NUMBER 4 751

Siphonosoma (Spengel, 1912)

Definition (revised).—Species usually of large size (greater than 5 cm in length
as adults) with introvert much shorter than trunk. Introvert with papillae and
sometimes also hooks arranged in rings. Body wall with coelomic extensions
(sacs); longitudinal muscle layer gathered into bands. Oral disk bears tentacles
arranged around the mouth (nuchal tentacles lacking). Four retractor muscles.
Contractile vessel with or without villi. Spindle muscle attached posteriorly
(sometimes with three roots anteriorly). Two nephridia. A single taxon no longer
separated into subgenera.

Critical Review of Species
Siphonosoma (Dasmosiphon) carolinense Fischer, 1928

Siphonosoma carolinense Fischer, 1928:138—140.—Nishikawa, 1977:11, fig. i
Siphonosoma hataii Sato, 1935:305—-308, pl. 3, fig. 7, text-figs. 3-7; 1939:373.

Material examined.—Sendai: Sato’s type of S. hataii (Palau 3)

Fischer’s material cannot be located but in Cutler and Cutler (1979:946—-948)
this species was relegated to the status of a junior synonym of S. cumanense.
The presumed difference in number of tentacles was demonstrated to be a false
dichotomy. We reaffirm that earlier conclusion.

Siphonosoma (Dasmosiphon) cumanense (Keferstein, 1867)

(See Stephen and Edmonds (1972) for synonymy prior to 1965)

Murina, 1967:1338; 1981:14—15.—Christie and Cutler, 1974:109.—Cutler, 1977:
138, fig. 1—Gibbs, 1978:85.—Cutler and Cutler, 1979:946-949.—Edmonds,
1980: 14—15, fig. 28.

Material examined.—Amsterdam: Sluiter’s type of Sipunculus claviger (V. Si.
168/6). Berlin: Grube’s type of S. cumanense var. semirugosus, Selenka et al.’s
types of S. cumanense var. vitreus and opacus. Hamburg: S. cumanense opacus
(V2059) & vitreus (V2051) determined by Fischer. Sendai: Sato’s types of S.
formosum (#6), S. koreae (#13), and S. hataii (Palau 3). Several hundred recently
collected specimens from the Indian and Pacific Oceans.

This common and morphologically plastic species has a long and convoluted
history in sipunculan literature. The most recent analysis of its variability is in
Cutler and Cutler (1979:946—949).

Based on recent Japanese collections Cutler, Cutler, and Nishikawa (in press)
conclude that it is conspecific with S. edule. The core of the argument is that as
a worm grows its introvert increases in length more slowly than the trunk so that
in large worms the introvert becomes proportionally shorter; S. edule is at one
end of a continuum but not a separate entity. Rather than submerge the more
common junior synonym, however, Cutler (in press) has petitioned the ICZN to
conserve S$. cumanense and suppress S. edule.

Siphonosoma (Dasmosiphon) edule (Pallas, 1774)

Lumbricus edulis Pallas, 1774:10-12.
Sipunculus edulis.—Sluiter, 1882:148—150, pl. 1, figs. 1 and 10; pl. 2, fig. 4, pl. 3,

752 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

fig. 1; 1886:484; 1891:122; 1902:5.—Selenka & deMan, 1883: 107—108 Tee
1899: 157—158.—Lanchester, 1905:29.
Siphonosoma edule.—Sato, 1939:371—373, pl. 20, fig. 6—Halder, 1975:58.
Sipunculus billitonensis Sluiter, 1886:487—488, pl. 3, figs. 1-2; 1891:123; 1902:4—
5.—Shipley, 1899:157, pl. 18, figs. 6-7; 1902:135.—Lanchester, 1905:30.
Siphonosoma billitonense.—Stephen, 1941:402.

Material examined.—Edinburgh: Stephen’s specimens from Maldive Islands
(1958.23.104). Amsterdam: Sluiter’s type of S. billitonense (V. Si. 168/1).

The status of this species has been examined elsewhere (Cutler and Cutler
1979; Cutler, Cutler, and Nishikawa, in press) and here we restate that it is
conspecific with S. cumanense. The presumed difference (a shorter introvert) has
been demonstrated to be attributable to large size and allometric growth. A pe-
tition to suppress this senior synonym and conserve S$. cumanense has been filed
with the ICZN (Cutler, in press).

Siphonosoma (Hesperosiphon) marchadi Stephen, 1960
Siphonosoma marchadi Stephen, 1960:515—516.

Material examined.—Paris: Stephen’s type-material (V-33).

This species was described on the basis of two incomplete specimens. Stephen
made a critical error by confusing the rectum with the esophagus. When the type-
material was examined it became evident that ‘““Rectum with numerous long villi’’
is really the contractile vessel with villi and ““The anterior portion of the esoph-
agus bears a large diverticulum”’ is rather the rectal caecum.

Therefore, this is not related to S. vastum as was asserted but in all ways is
comparable to S. cumanense. We propose that the name S$. marchadi be sub-
merged as a junior synonym of S. cumanense.

Siphonosoma (Hesperosiphon) parvum Fischer, 1928
Siphonosoma parvum Fischer, 1928:141-143, figs. 1-2.

Material examined.—None.

This single worm could not be located and is assumed to be lost. Its uniqueness
has been questioned but no formal action proposed. Its separate status rests on
a presumed difference from S. vastum in the size and number of rectal caecae.
No actual counts or measurements were given; it was stated only that they were
larger and less numerous in S. parvum. Our experience shows that such structures
do exhibit variation within a population and this seems a clear case of an artificial
dichotomy within a geographically contiguous population. Therefore, we herein
consider S$. parvum to be a junior synonym of S. vastum.

Siphonosoma (Hesperosiphon) vastum
(Selenka, De Man and Bulow, 1883)

Sipunculus vastus (Selenka et al., 1883:103-104, pl. 12, fig. 171, pl. 13, fig. 179.—
Shipley, 1898:469; 1899: 158; 1902:136.—Augener, 1903:315—317.—Fischer, 1919:
279-927-2200:

Siphonosoma. vastus. —Wesenberg-Lund, 1937:2-5, figs. 1-2.

VOLUME 95, NUMBER 4 53

Siphonosoma vastum.—Edmonds, 1955:92—95, figs. 8-9.—Cutler & Cutler, 1979:
949.—Edmonds, 1980: 15-16, fig. 31.—Murina, 1981: 15—16.
Siphonosoma crassum Spengel in Fischer, 1919:279; 1927:199.

Material examined.—Berlin: Selenka’s type #918, 2 spec. from Jaluit, Marshall
Islands; from Mozambique #279, | spec. Copenhagen: Wesenberg-Lund’s 2 spec-
imens from Indonesia.

The two type-specimens have dried out but the third worm is in good condition
and matches the description. As pointed out in the section on S$. parvum, we
consider these two species to be conspecific. As this is the older name, it remains
the valid senior synonym.

Siphonosoma (Siphonosoma) amamiense (Ikeda, 1904)

Sipunculus amamiense Ikeda, 1904:36—38, figs. 64-65; 1924:31.
Siphonosoma amamiense.—Sato, 1939:371.—Cutler and Cutler, 1981:57-58.

Material examined.—Tokyo: Ikeda’s type-material. Sendai: Specimens identi-
fied by Sato (#3-—12). One recently collected specimen from Yaeyama Islands.

When Ikeda described this species he compared it to S. funafuti and said, “*. . .
S. funafuti is so peculiar in color and body form that it bears no resemblance to
the present species in external aspects.’’ He also noted the absence of fixing
_muscles and the peculiar form of the papillae in S. funafuti. In the section on S.
funafuti we comment on the fragile condition of those worms and dubious quality
of fixing muscles as a distinguishing character. In Cutler and Cutler (1981) we
comment on some morphological aspects of Ikeda’s worms, e.g. true contractile
vessel villi are not present, and draw attention to the similarity with S. funafuti.

Despite Ikeda’s thought that these species differ in external color and form
(see Cutler and Cutler 1981:53-55 for more comments on this point), we conclude
that they are conspecific and despite the fact that Ikeda’s description is a better
one, must regrettably submerge S. amamiense as ajunior synonym of S. funafuti.

Siphonosoma (Siphonosoma) arcassonense (Cuenot, 1902)

Sipunculus arcassonense Cuenot, 1902:15, pl. 1, figs. 15; 1922:1314, figs. 7a
and 7b.
Siphonosoma arcassonense.—Saiz Salinas, 1980:9092, fig. -24.

Material examined.—Two specimens recently collected from type-locality: Ar-
cachon, France. Nancy: Undissected specimen from Cuenot’s material.

Cuenot’s type had been reported to be in the Paris Museum but was located
at the Université de Nancy, France. In his unpublished Master’s thesis, Saiz
Salinas described several more specimens from the type-locality. He stated that
his worms have four fusiform bodies at the extreme posterior end of the trunk
but are difficult to see because of the coagulation of the coelomocytes. He also
said contractile vessel villi are present but does not describe them in any detail.

The 240 mm paratype (?), a 150-mm specimen Saiz Salinas loaned us, and a
135-mm worm sent by C. Cazaux, have very many, crowded, well-developed
digitiform or shorter clavate villi arranged along most of the contractile vessel.
The three or four, thin, pale tubular fusiform bodies are about 4-7 mm long and

754 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

more delicate than those in S. ingens. Each of the specimens we examined has
the posterior tip of the trunk drawn in by 2—3 mm and the ventral nerve cord
gives off numerous large, transverse nerves forming a plexus near the posterior
end. The scalelike ‘hooks’ are arranged in up to 120 rings and are strikingly similar
to S. dayi, i.e., tubular, attached to the skin except for the very tip and lacking
a sharp point (Fig. 1A). This active burrower occurs sympatrically with Sipun-
culus nudus on beaches at Arcachon, France, but is rare.

We do not propose any change in the status of this peculiar and localized
species.

Siphonosom (Siphonosoma) australe (Keferstein, 1865)

Siphonosoma australe.—Halder, 1976:3.—Cutler, 1977:139.—Cutler and Cutler,
1979:949. (See Stephen and Edmonds (1972) for synonymy prior to 1965).

Material examined.—London: Baird’s type of S. aeneus. Sendai: Sato’s type
of S. takatsukii (Yap 6). Numerous specimens recently collected from the Indian
Ocean.

Keferstein’s specimens cannot be located but this is a well known, widely
reported, and valid species.

Siphonosoma (Siphonosoma) boholense
(Selenka, De Man and Bulow, 1883)

Sipunculus boholensis Selenka et al., 1883:109-111, pl. 12, figs. 175—177.—Fisch-
er, 1895:5—6.—Lanchester, 1905b:27.
Siphonosoma boholense.—Edmonds, 1980:17.

Material examined.—Berlin: Selenka’s type (#988). Hamburg: Fischer’s spec-
imen from Bohol (V 2050).

This species has been described in recent years as having contractile vessel
villi, e.g. “*. . . with numerous very small villi, appears bushy’’ (Edmonds 1980:
17). Stephen and Edmonds’ (1972:63) translation of Selenka’s description says,
**.. with many tubules.’ Selenka’s words (1883:110) are ‘‘mit zahlreichen
braunrothen Blindsackchen besetzt.’’ The term commonly used in German for
villi is “zotten’ and the Blindsackchen which translates as ‘little caecae,’ suggests
something different. As discussed in the introduction, there has been confusion
here. True villi are digitiform, longer than broad and quite distinct, well-defined
structures. However, in some worms lacking villi, the contractile vessel is large
and exhibits bulbous vesicular or bubble-like swellings on the surface. When we
examined Selenka’s types we saw only a bulbous or pleated vessel. Fischer’s
worm in Hamburg shows no villi (he made no reference to any in his paper).
Stephen and Edmonds (1972:63) reported re-examining two specimens from North
Borneo in the British Museum. They made no reference to villi; we saw none
either.

So, if this species lacks contractile vessel villi and hooks and has its retractor
origins at different levels, it is similar to S. amamiense and S. funafuti. These
two species are said to have fewer than 20 longitudinal muscle bands while S.
boholense has 29-33. We would point out that only 10 specimens have been
named S. boholense in the last 100 years and that none of these is less than 200

VOLUME 95, NUMBER 4 WD

mm long. This raises the question: are these worms a distinct species or only an
older, larger subset of a population which also includes S$. amamiense and S.
funafuti? In other words, do small S. boholense really have 29-33 longitudinal
muscle bands or have these bands split as the size increased, in a few older
worms?

We seem to have in these three species two ends of a continuum with organisms
having trunk lengths between 120 and 200 mm unrepresented in the collections.
While we are inclined to consider these taxa as conspecific, we will postpone
such a decision. We anticipate, however, that intermediate sized specimens can
be found which will have 20-30 longitudinal muscle bands and thereby connect
these two subsets. Also, histological and biochemical work may solve our un-
certainty.

Siphonosoma (Siphonosoma) bonhourei (Herubel, 1904)
Sipunculus bonhourei Herubel, 1904:479—480; 1907:137-141, figs. 30-34.

Material examined.—None.

This species is known from only a single specimen which cannot be located.
Herubel’s description is long and detailed but it is unclear whether or not he had
an entire worm. The origin of retractor muscles at the beginning of the posterior
third of the trunk and the ratio of introvert to trunk (80 mm to 140 mm trunk)

'make one suspicious. Herubel also says that the introvert is 1/7 the trunk length.

There are other unclear issues such as the contractile vessel with short lateral
ramifications or digitations (Herubel does not illustrate villi). This is in the com-
plex which includes S. amamiense, S. funafuti, and S. joubini. In view of the
uncertainties and loss of the type-specimen, it seems most prudent to place this
species in incertae sedis.

Siphonosoma (Siphonosoma) dayi Stephen, 1942

Siphonosoma dayi Stephen, 1942:246-247, pl. 11, figs. 1-2.—Wesenberg-Lund,
1963:103, fig. 1.

Material examined.—Edinburgh: Stephen’s type #1958.23.1.

Only three specimens have been given this name (one by Stephen and two by
Wesenberg-Lund) but it does seem to be unique, if rare and geographically re-
stricted. The unique feature is its introvert papillae which are well illustrated by
both authors and referred to as scalelike bodies/structures or simply scales. One
fact not mentioned is that the entire unit is fixed to the skin, not only basally as
is a hook. There also appears to be an open pore at one end. It is easy to suppose
that these are homologous to hooks and that their development became modified
in some way. It may well be that we see here an example of an intermediate stage
in the development of the S. australe type of hooks or perhaps a neotenous form.
Siphonosoma australe is a common sympatric species so these three worms may
simply be a few anomolous individuals. More data are needed to determine whether
or not these really represent a biological population.

For the present we propose no change in status but point to its similarity to S.
australe.

756 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Siphonosoma (Siphonosoma) eniwetoki Fisher, 1950
Siphonosoma eniwetoki Fisher, 1950:805-808, pl. 1.

Material examined.—Washington, D.C.: Fisher’s type #21128, plus bottle of
7 specimens (#24603) from Likiep Atoll, Marshall Islands; 12/13/51; identified by
Fisher but unpublished.

These worms have distinctive introvert hooks. As drawn by Fisher they are
closely associated with large papillae, but when the skin is stretched as in an
expanded worm, the height of the papillae decreases. The apparent shape of the
hook is dependent upon the angle at which one views the hooks (Fig. 1D). The
sharpness or bluntness of the point can be misleading. The contractile vessel villi
are digitiform or clavate but distinct even if small. They seem to be present along
only the free part of the esophagus.

When Fisher described this species, S. rotumanum was still thought to lack
hooks. This error was corrected by Edmonds (1971). As a result, Fisher did not
compare his worms with S. rotumanum. But, with our correct understanding of
this taxon, a comparison of these forms shows them to be conspecific. As S.
rotumanum (Shipley, 1898) is the older name, S. eniwetoki becomes a junior
synonym of that species.

Siphonosoma (Siphonosoma) funafuti (Shipley, 1898)
Sipunculus funafuti Shipley, 1898:469, pl. 37, figs. 4-5.

Material examined.—London: Shipley’s paratype, | specimen. Hamburg: Ship-
ley’s type (?), 4 specimens.

These five worms all seem to have been identified by Shipley but the issue of
holotype/paratype/cotypes is unclear. As noted by Stephen and Edmonds (1972:
65), ‘‘The body wall is thin and the internal structures very fragile.’’ None of
these has villi on its contractile vessel. The one in London does exhibit bulbous
vesicles which have been interpreted as villi by some biologists. The alleged
difference noted by Stephen and Edmonds (1972) between this species and S.
amamiense is the number of intestinal fixing muscles (0 vs. 2). In previously
dissected material it is impossible to say with certainty that such fragile structures
were never present. The taxonomic significance of this character is doubtful even
if it were actually absent and not just overlooked or inadvertently broken.

We conclude that S. funafuti is a valid species, conspecific with, and a senior
synonym of S$. amamiense.

Siphonosoma (Siphonosoma) ingens (Fisher, 1947)

Siphonomecus ingens Fisher, 1947:365—368, pl. 14, 15.
Siphonosoma ingens Fisher, 1952:382—385, pl. 20, 21.

Material examined.—Washington, D.C.: Fisher’s type (#20910).

This species with fusiform bodies, contractile vessel villi and without introvert
hooks remains a valid species. It is very similar to S. mourense (Sato) except for
the presence of the peculiar fusiform bodies (Fig. 11). Preliminary histological
examination of these bodies suggests an excretory function.

VOLUME 95, NUMBER 4 WS

Siphonosoma (Siphonosoma) joubini (Herubel, 1905)
Sipunculus joubini Herubel, 1905:51—54, 3 figs; 1907:141-145, figs. 35-37.

Material examined.—None.

The single worm Herubel described cannot be located and no one else has
reported such an animal in the intervening 77 years. Herubel’s (1907) Fig. 37
shows a rather unique contractile vessel with a few small caecum-like structures
(digitations), not normal villi. Whether this animal represents a population or
simply a single anomolous individual is uncertain but the fact that in the many
Indo-West Pacific collections, no other specimens have appeared, makes the
latter case probable.

In view of the above facts we propose placing this species on the list of incertae
sedis.

Siphonosoma (Siphonosoma) mourense Sato, 1930

Siphonosoma mourense Sato, 1930:6—-8, pl. 1, figs. 2-4; 1937:148-149, pl. 3, fig.
9; 1939:370-371.—Leroy, 1936:4235.

Material examined.—Sendai: Sato’s type-material (#183, 3 specimens) plus
others identified by Sato. )

This species was discussed in Cutler and Cutler (1981) and at that time consid-
_ ered to be a valid species. It is similar to S. cumanense except that the retractor
muscles originate at different levels and there are no transverse dissepiments. It
is also very similar to S. ingens except that this latter species has the peculiar
and easily overlooked fusiform bodies at the posterior end of the trunk. An ex-
amination of Sato’s worms did not show fusiform bodies to be present. Siphon-
osoma moursense and S. ingens are closely related, both occurring in the North
Pacific.

Siphonosoma (Siphonosoma) novaepommeraniae (Fischer, 1926)

Sipunculus novaepommeraniae Fischer, 1926:104—106, pl. 3, figs. 2-4, 6.
Siphonosoma novaepommeraniae.—Wesenberg-Lund, 1959:55-58, text-figs. I-
3.—Murina, 1967: 1338.—Edmonds, 1971:140-141; 1980:17.—Cutler, 1977:139.

Material examined.—Hamburg: Fischer’s type (#V3414). Copenhagen: Wes-
enberg-Lund’s specimens from Mauritius.

In Cutler and Cutler (1979:948) in an analysis of transverse dissepiments in
Siphonosoma cumanense, it was observed that ‘*. . . S. novaepommeraniae is
exactly like S. cumanense except it lacks the transverse dissepiments. Wesen-
berg-Lund (1959) says they exist in her Mauritius forms of S$. novaepommeraniae
and even draws them. Hers then should be considered S. cumanense and possibly
Fischer’s (1926) should be here also.”’

Our examination of Wesenberg-Lund’s worms reinforced this conclusion.
Fischer’s single worm did not exhibit these structures but as shown in Cutler and
Cutler (1979) this character is not consistently present in a population and should
not be weighed very heavily. Like Fischer, the records of Murina (1967) and
Edmonds (1980) are single individuals. It is our conclusion that S$. cumanense is

758 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

a common, circumtropical species which includes some specimens with and some
without transverse dissepiments and that when certain careful observers noticed
their absence, a different name was applied. However, we propose that this
distinction is taxonomically meaningless and S. novaepommeraniae should be
treated as a junior synonym of S. cumanense.

Siphonosoma (Siphonosoma) pellucidum (Sluiter, 1902)
Sipunculus pellucidus Sluiter, 1902:9-10, pl. 1, fig. 3.

Material examined.—Amsterdam: Sluiter’s type (V.Si. 168.7), 2 spec.; (V.Si.
168.8), 3 spec.

These five worms range in trunk length from 12-20 mm. They were not shifted
from Sipunculus to Siphonosoma until Stephen and Edmonds (1972): “‘A re-
examination of one of Sluiter’s specimens (not marked as type) shows that the
species 1s a Siphonosoma. ... The introvert is not strongly marked from the
trunk, it lacks the triangular, squamiform papillae of a Sipunculus and there are
no hooks or spines.”’

The small size clearly shows these to be juvenile worms, as does their semi-
transparent, thin body wall. They appeared to us to strongly resemble young
Sipunculus species. The development of muscle bands, both longitudinal and
circular as well as introvert papillae may well occur later in their ontogeny. If
these are truly Siphonosoma, they are the only ones lacking villi and hooks, and
which have the retractor origins at the same level.

We propose that one cannot make a definite assignment on the basis of this
material but favor the original placement in the genus Sipunculus. However, due
to their condition and small size, it seems best to place this taxon in species
inquirendum pending future clarification.

Siphonosoma (Siphonosoma) pescadolense Sato, 1939
Siphonosoma pescadolense Sato, 1939:376—379, pl. 20, fig. 8, text-figs. 18-22.

Material examined.—None.

Cutler and Cutler (1981) synonymized this species with Sato’s species, S. tak-
atsukii, and made the latter a subspecies of Siphonosoma australe. We reaffirm
that action.

Siphonosoma (Siphonosoma) rotumanum (Shipley, 1898)

Sipunculus rotumanus Shipley, 1898:469-470, pl. 37, figs. 1-3.

Siphonosoma rotumanum.—Edmonds, 1971:143—144; 1980:18; Christie & Cutler,
1974: 109-110.—Gibbs, 1978:85.

Siphonosoma hawaiense Edmonds, 1966:386—388, figs. 1—4.

Material examined.—London: 1 specimen identified by Shipley, possible type.
Hamburg: | specimen identified by Shipley; possible cotype (V5391).

When Shipley described this species he misinterpreted the unique introvert
hooks/spines which are present by referring to them as papillae. This oversight
which created problems for Edmonds (1971) also caused Fisher (1950) to bypass
this species when he named S. eniwetoki (as noted earlier in this paper). The

VOLUME 95, NUMBER 4 159

peculiar introvert hooks and associated papillae can vary somewhat in apparent
form in different states of contraction or when viewed from different angles. This
species name remains valid as redescribed by Edmonds.

Siphonosoma (Siphonosoma) takatsukii Sato, 1935

Siphonosoma takatsukii Sato, 1935:308—310, pl. 3, fig. 8, text-figs. 8-10; 1939:
BSE

Material examined.—Sendai: Sato’s type-specimen.

Cutler and Cutler (1981) reduced this to a subspecies of Siphonosoma australe.
There is only one specimen and it differs from S. australe in having smaller,
blunter hooks, and being restricted to the northeast Pacific Ocean. That conclu-
sion is reaffirmed.

Known Distribution of Siphonosoma Species

Only three of these ten species have been recorded from a broad area; the
remaining seven apparently have a much more restricted range as follows:

S. arcassonense Western France

S. ingens Southern California

S. mourense Northeast Honshu, Japan

S. dayi Natal, South Africa

S. boholense Queensland, Australia, and North Borneo

S. funafuti Southern Japanese Islands, and Funafuti

S. rotumanum Hawaii to Queensland, and Cape Province, South Africa
S. vastum Indo-West Pacific—tropical

S. australe Indo-West Pacific—tropical and subtropical

S. cumanense All oceans, tropical and subtropical (not recorded from the

eastern Atlantic and eastern Pacific Oceans)

The first two species have a very restricted distribution, are separated by thou-
sands of miles from other Siphonosoma, and are in the only regions unoccupied
by S. cumanense. They are also the only two with posterior fusiform bodies.

Key to Siphonosoma Species

1. Contractile vessel without distinct villi (bulbous vesicles may be present)

Pet ees Menren OTe LAM Ti MONE Cae bee TaiAs. 2 LPtiGhS OM BOOS « ucteneet 2
=w@anmuacile vessel bears distinct villi 2.00) ic Se a eee 6
Pemliinevienrt with Wooks-or scalelike papillae ............-.-...+....-.4..- 3
=walntcovert without hooks or scalelike papillae .....2.........-.-.-.000.. 5
3. Introvert with rings of tubular scalelike papillae, not free standing hooks
ne fe tee CN FT SM AROS, © MND AN ey ho lie al hoes 0 te aos al S. dayi
MEOW ETL WitMGIStinNcl MOOKS 3. oe. so ek el, BE LS. PI 4
4 Rectum with numerous caecae/diverticula .....................- S. vastum
— Rectum without numerous caecae/diverticula ................. S. australe
Sealeessuihan 22 loneitudinalimuscleibands) 252255 -25.505.05---. - S. funafuti

=~evionre tnan 2S longitudinal muscle bands) ~5....:-...-22- 5: S. boholense

760 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

6. Introvert with rings of hooks or scalelike papillae ...................... qi
= Introvert without hooks’ or scalelike*papillae™= 25.1.4 ee eee 8
7. Scalelike chitinized papillae on introvert; posterior end of trunk with in-
ternal ‘fusiforme bodies 7,5) eae, eee ees eee nae ne S. arcassonense
— Short, blunt hooks on introvert; no fusiform bodies ........ S. rotumanum
Sue Retractor muscles oneinate ausameleveleres . suerte S. cumanense
— Dorsal retractor muscles originate anterior to the ventral pair ........... 2
9, Pustionm: bodies inspostenonm end omminks. 600 oe eee S. ingens
Fusiform bodiessabsemt ti. 2 5 ee en eee S. mourense
Acknowledgments

This work was made possible with the cooperation of the following persons
who made specimens available to us: H. Terayama, University Zoological Mu-
seum, Tokyo; Z. Kawabata, Tohoku University, Zoological Institute, Sendai; S.
van der Spoel, Instituut voor Taxonomische Zoologie, Amsterdam; J. Renaud
Mornant, Museum National d’Histoire Naturelle, Paris; G. Hartwich, Museum
fur Naturkunde, Berlin, DDR; M. Dzwillo, Universitat Zoologisches Museum,
Hamburg; J. B. Kirkegaard, University Zoological Museum, Copenhagen; S.
Chambers, Royal Scottish Museum, Edinburgh; R. W. Sims, British Museum,
(Natural History), London; M. Jones, National Museum of Natural History,
Washington, D.C.; J. Saiz Salinas, Bilboa, Spain; C. Cazaux, Station Biologique
D’ Arcachon, France; B. Conde, Université de Nancy, France. The helpful com-
ments and suggestions of P. Gibbs, Plymouth, are gratefully acknowledged. A
draft of this paper was read by T. Nishikawa (Nagoya) and S. Edmonds (Ade-
laide). This work was supported by a grant from the U.S. National Science Foun-
dation (DEB 8011121).

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Edmonds, S. J. 1955. Australian Sipunculoidea. I. The genera Sipunculus, Xenosiphon, and Si-
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VOLUME 95, NUMBER 4 761

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762 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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Stephen, A. C., and S. J. Edmonds. 1972. The Phyla Sipuncula and Echiura.—London: British
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Utica College of Syracuse University, Utica, New York 13502, U.S.A.

PROC. BIOL. SOC. WASH.
95(4), 1982, pp. 763-765

A NEW SPECIES OF EUGLANDINA FROM PERU
(GASTROPODA: PULMONATA: SPIRAXIDAE)

Fred G. Thompson

Abstract.—Euglandina haasi is described from the Maranon Basin. It is most
closely related to E. cylindrus, which also occurs in the Maranon Basin. Euglan-
dina haasi is the fourth species of Euglandina recorded from Peru. Euglandina
c. cylindrus (Martens) originally was described from ‘*‘Maranon-Flusse, Colom-
bia.’ This is corrected to Maranon River Basin, Peru.

Carnivorous land snails of the genus Euglandina are sparcely represented in
South America. The genus underwent its principal radiation in Mexico, and only
a few species-groups have migrated southward as far as the northern Andes. Four
species are known from Peru. Haas (1952:117) and Weyrauch (1958: 105) discuss
the occurrence in Peru of the widely distributed E. striata (Muller). Weyrauch
(1960:25—26) describes E. altispira from the Maranon Basin, Depto. Cajamarca.
The species is closely related to E. striata. Haas (1951:539-540) describes E.
cylindrus augusta. This subspecies also occurs in the Maranon Basin, Depto.
‘Cajamarca. The fourth species is described here. It is related to E. c. augusta
and comes from a nearby area in the Maranon Basin.

Euglandina haasi, new species
Figs. 1—4

Type-locality. PERU, Depto. Cajamarca, 5 km WSW of Balsas, 1220 m alti-
tude. Balsas is a small village on the Rio Maranon in the Depto. Amazonas and
is 54 km ENE of Cajamarca. Holotype: Florida State Museum, UF 36122; col-
lected 24 April 1972 by Fred G. Thompson, under a limestone boulder. The
Maranon Basin at this locality is arid with sparce growths of columnar cactus
and thorn scrub.

Description.—Shell medium-sized, cylindrico-terete (Figs. 1-2), thin but solid,
translucent, glossy, corneous-brown with occasional varix-like whitish streaks.
Spire slender; nearly straight-sided; longer than aperture. Holotype with 7.8 whorls.
Embryonic shell (Fig. 3) with 3.8 whorls bearing very shallow suture. First whorl
slightly raised above the following, forming a rounded dome. Embryonic whorls
smooth, sculptured along suture with occasional weak vertical striations. Follow-
ing whorls arched and with rather distinct shoulder. Suture deeply impressed.
Body whorl nearly cylindrical. Postembryonic whorls sculptured with weak growth
striations forming poorly defined thread-riblets; most distinct on shoulder of whorls,
suture crenulate (Fig. 4). Weak but distinct subsutural impressed spiral line caus-
ing crenulations to be beaded. Aperture elongate-auriculate, about 0.46 times
length of shell. Middle of outer lip strongly arched forward (Fig. 2). Columella
truncate, obliquely twisted to the right (Fig. 1) and forward (Fig. 2).

Length, 46.8 mm; width, 16.1 mm; aperture height, 21.7 mm.

Remarks.—Euglandina haasi is most closely related to E. c. cylindrus (Mar-

764 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-4. Euglandina haasi, Holotype: |, Frontal view; 2, Lateral view; 3, Apex; 4, Sculpture
along suture of last whorl on posterior side.

tens) and E. c. augusta Haas. The three are alike in the shape of the embryonic
whorls, size, general shape and whorl count. They also occur in the same area
of Peru (see below). Euglandina haasi differs from the two subspecies of E.
cylindrus by having a distinct subsutural impressed spiral line beading the cren-

VOLUME 95, NUMBER 4 765

ulations, a smoother shell with finer incremental striations, a twisted columella,
shouldered whorls and a deeply impressed suture. Euglandina cylindrus has a
weakly crenulate suture that lacks a subsutural spiral line, the shell is densely
sculptured with rib-striations; the columella is straight and vertical, the whorls
are weakly arched and lack shoulders, and the suture is shallow. Complete syn-
onymies for the two subspecies of E. cylindrus are given below.

Etymology.—I take pleasure in naming the species for the late Dr. Fritz Haas,
(1886-1969), who contributed greatly to our knowledge of Peruvian mollusks.

Euglandina cylindrus cylindrus (Martens)

Glandina cylindrus Martens, 1860, in Albers:29.—Strebel, 1875:22—23, Taf. I, fig.
39 (embryonic whorls), Taf. X, figs. 39-39b (shell).

Oleacina cylindrus (Martens).—Pfeiffer, 1868:279.—Tryon, 1885:34, pl. 8, fig. 8
(shell).

Euglandina cylindrus (Martens).—Pilsbry, 1907: 184.

TypeHocality.—Martens (in Albers, 1860:29) describes this species from the
‘*Maranon-Flusse, Colombia.’’ Subsequent authors interpreted this to be the
‘‘Maranon River, Colombia’’ or the ‘‘upper Andes, Colombia.’’ There is little
doubt that the correct geographic origin of the type is the Maranon River Basin,
Peru, which places it in close geographic proximity to its most closely related
_allies. The assignment of E. c. cylindrus to the Colombian fauna was erroneous,
as was typical of many species of land snails described during the nineteenth
century. It is known only from the holotype in the Berlin Museum.

Euglandina cylindrus augusta Haas
Euglandina cylindrus augustua Haas, 1951:539-541, fig. 122 (shell).

Type Hocality.—PERU, Depto. Cajamarca, Jaén, 700 m alt. Holotype: Field
Museum of Natural History 30911; collected by Wolfgang K. Weyrauch.

Literature Cited

Albers, J. C. 1860. Die Heliceen nach nattrlicher Vervandtschaft systematisch geordnet. II. Aus-
gabe, herausgegeben von E. von Martens: 1-359. Leipzig.

Haas, F. 1951. Remarks on and descriptions of South American non-marine shells —Fieldiana 31:

503-545; figs. 97-126.

1952. South American non-marine shells: further remarks and descriptions.—Fieldiana 34:

107-132, figs. 14-26.

Pfeiffer, L. 1868. Monographia Heliceorum Viventium, VI. 1-598. Leipzig.

Strebel, H. 1875. Beitrag zur Kenntniss der Fauna Mexikanischer Landund Stisswasser-conchylien,
II. 1-58; Tafs. I-XIII. Hamburg.

Tryon, G. W. 1885. Manual of Conchology, second ser., 1:1—364; pls. 1-60. Philadelphia.

Weyrauch, W. K. 1958. Neue Landschnecken und neue Synonyme aus Sudamerika, 1.—Archiv

fur Molluskenkunde 87:91—139, Tafs. 6—9a, figs. 1-51.

. 1960. Zwanzig neue Landschnecken aus Peru.—Archiv ftir Molluskenkunde 89:23—-48, Tafs.

3-6, figs. 1-44.

Florida State Museum, University of Florida, Gainesville, Florida 32611.

PROC. BIOL. SOC. WASH.
95(4), 1982, pp. 766-771

DESCRIPTION OF A NEW CYPRIDOPSINE GENUS
(CRUSTACEA: OSTRACODA) FROM CAMPBELL
ISLAND, WITH A KEY TO THE CYPRIDOPSINAE

K. G. McKenzie

Abstract.—Austrocypridopsis, a new genus of cypridopsine ostracodes with
the type-species Austrocypridopsis terrilli n.sp. is described from Campbell Is-
land. A key to Cypridosinae is provided, and the subfamily Pseudocyprettinae is
designated as new.

As shown by Smith and Sayers (1971, Table II) no Ostracoda have been re-
corded previously from Campbell Island; and only one ostracode species Cypretta
sp. has been recorded from nearby Macquarie Island (Lofthouse 1967). Never-
theless, some eight species of continental Ostracoda are known from the Antarctic
mainland and from sub-Antarctic islands. Of these eight species, only two occur
at more than a single locality: Ilyodromus kerguelensis Muller, 1908, is recorded
from four sub-Antarctic islands in the Southern Ocean; and Notiocypridopsis
frigogena (Graf, 1931) occurs on South Georgia and Signy Island (Smith and
Sayers, 1971). The present record increases this number to nine species and
necessitates the description of a new cypridopsine genus.

Material

The material was collected by Mr. P. Terrill, of the Australian Museum, on a
saddle between Mount Azimuth and Mount Fizean, Campbell Island, among plants
and sediment from the headwaters of a small stream flowing north, at an altitude
of 400 m. The collection date was 6 January 1980.

A total of 9 specimens was collected, comprising 5 adult females, 1 juvenile
(A-1) female, | empty female left valve, 1 adult male and | juvenile (A-1) male.
All the specimens have been deposited in the Australian Museum, some as dis-
section slides. The Accession Number of this collection is P 30226; Holotype
(male) is AM P31528; Paratype (female) is AM P 31527; remaining Paratypes are
AM P30266.

Cypridopsidae Kaufmann 1900
Cypridopsinae Kaufmann 1900
Austrocypridopsis, new genus

Etymology.—A combination of auster (L.) = south, and the generic suffix—
Cypridopsis.

Diagnosis.—Resembling other cypridopsines in that both segments of the max-
illule palp are elongate but differing from them in: the subrectangular carapace
shape (in lateral view); the possession of reduced ‘‘natatory”’ bristles on the
antennae; the absence of an epipod on the maxilla (P1); and in the relatively
powertul (for a cypridopsine) furca. Both sexes known.

Type-species.—Austrocypridopsis terrilli, new species.

VOLUME 95, NUMBER 4 767

Discussion.—At first sight this genus appears to be a candonid because of its
subrectangular shape which is unusual among cypridopsines. However, when the
valves are examined in detail, the muscle scars are observed to have the char-
acteristic cypridopsid pattern. Further, the furca, while unusually well developed
for a cypridopsine, is definitely non-candonid. Finally, candonids are character-
ised by the absence of ‘“‘natatory’’ bristles on the antennae, while in the new
genus reduced ‘‘natatory”’ bristles are clearly present.

In its subrectangular shape, the new genus also resembles the genus Callisto-
cypris Schornikov, 1980, described by its author as the type-genus of a new
cypridopsid subfamily. But, the adductor muscle scar pattern of Callistocypri-
dinae is unique among cypridopsids (Schornikov 1980, Figs. 1, 2), consisting of
six scars arranged biserially (four anterior, two posterior) whereas in Austrocy-
pridopsis the adductor muscle scars are four in number (three anterior and one
(divided) posterior). Further, the two mandibular scars are proportionately larger
in Austrocypridopsis than in the two known genera of Callistocypridinae (R. H.
Victor personal communication, July 1981, has a second callistocypridine genus,
from container habitats in the Pacific islands).

The Diagnosis indicates the characters which distinguish this genus from other
cypridopsine genera. Apart from the subrectangular shape, which is distinctive
enough to isolate this genus immediately from Cypridopsis (subovate), Sarscy-
pridopsis and Plesiocypridopsis (subquadrate), Cavernocypris (subtriangular,
Notiocypridopsis (subtrapezoid), and Kapcypridopsis (subreniform); the furca is
much more powerful, with a stronger terminal claw and broader proximal part
than in these genera. It shares the reduced “‘natatory’’ bristles of the antenna
with Cavernocypris but that genus is so adapted for existence in (subsurface)
cave systems whereas Austrocypridopsis is part of the surface benthos. Some
other cypridopsine genera are characterized by well-developed antennal natatory
bristles and are good swimmers. In any case, Austrocypridopsis differs markedly
from Cavernocypris because whereas a maxillar epipod is absent in the former it
is clearly present in Cavernocypris species although the number of Strahlen is
less than normal, varying from 2—S (Hartmann 1964; McKenzie 1972). Further,
the furca in Cavernocypris is either altogether lacking or much reduced (Hart-
mann 1964; McKenzie 1972) whereas it is exceptionally powerful in Austrocy-
pridopsis, which is thereby well adapted for crawling actively over the substrate.

Using these characters—subrectangular shape, reduced antennal “‘natatory”’
bristles, absent maxillar epipod, and relatively powerful furca—it is easy to key
out Austrocypridopsis from the previously described cypridopsine genera.

Key to Genera of the Subfamily Cypridopsinae’

1. Shell whitish to pale brown, ornamented with pits arranged concentrically
SeereeaeUiabe or Coucentically: ribbed. | .22sc 0. fletiiadvan- evils ec) 282 2
NMRA Si no Macs cata df Ss hb Se ne dR = BS nie 3

1 Three related living genera are not included. They are Oncocypris, Oncocypria and Pseudocy-
pretta. The first two are usually synonymized (Oncocypris is the senior synonym). Oncocypris has a
distinctive pustulose surface ornament, the inner lamella bears characteristic lunettes and the max-
illule 3rd lobe has 4 Zahnborsten. It well merits the separate subfamilial status already accorded to
it by modern researchers, viz. Oncocypridinae De Deckker 1979.

768 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

2. Shell whitish, ornamented with pits arranged concentrically, or tubercu-
late; right valve margin smooth; cleaning leg (P3) with terminal pincer
(Fanee) ide ee Sete Se OS: oR at Aa See natin a a Oa na Zonocypris

— Shell pale brown, concentrically ribbed; right valve margin denticulate
and bearing midventrally a small boss; cleaning leg (P3) simple terminally,

without a pincer (Zamee)™ ors. 04, Gene ae ee em pen eine Neozonocypris
3. Shell subovater 10... cs res ARE A ened EAT A ee 4
~ “Shelldifferemt soy ne ee OSE Atco er >
4. Right valve ventromarginally tuberculate and overlapping left valve ven-
Pray ee ee ee eee, IR ae ES OPO EE RRR Cypridopsis
— Right valve smooth ventromarginally, left valve overlapping right valve
ventrally 72 EA OS EE? Ce BIS Don hE oct een are tote STCR ay Neocypridopsis
5. Shell subreniform or subtrapezoid, inner lamellae broad anteriorly and
posteriorly, antennal natatory bristles reduced or well developed ........ 6
— Shell of different shape, inner lamellae broad anteriorly but narrow pos-
teriorly, antennal natatory bristles reduced or well developed ........... 7
6. Shell subreniform, antennal natatory bristles reduced ...... Kapcypridopsis
— Shell subtrapezoid, antennal natatory bristles well developed ..........
Seok a alt had CGE? MM MEI LALA dae tries S87 E Gene Biase ea! Notiocypridopsis
7. Shell subquadrate, antennal natatory setae well developed .............. 8
— Shell subtriangular or subrectangular, antennal natatory setae reduced ... 9
8. Male clasping palps elongate, hemipenis with anterior lobe upturned ...
ee Re TN eae NPR ee eT GR ee OP ee a SCTE 5 oe |e Plesiocyopridopsis
— Male clasping palps normal, hemipenis with anterior lobe downturned
polenta TER PARIS Ta Me Ae Sepa Se VES OOD, eS anseypnidopsys
9. Shell subtriangular, furca absent or reduced, maxillar epipod present (with
2=5°Strahlen)<¢ A ee es Pe CD, Paid RA Rs ee Soran Sata Cavernocypris
— Shell rectangular, furca powerful (for a cypridopsine), maxillar epipod
AWSEMO LR e. Ee REALE SO ee ene aah aien eee a) Austrocypridopsis

Austrocypridopsis terrilli, new species
Figs. 1-18

Etymology.—For Mr. P. Terrill who collected the material.

Description.—Shell whitish to cream-colored, medium-sized (length about 0.75
mm). In lateral view subrectangular, smooth, adorned with scattered fine hairs;
anterior broadly rounded; dorsum relatively straight and sloping backwards; pos-
terior broadly subtruncate; venter inflexed medially; right valve slightly larger
than left valve; greatest height anteromedial and about half the length. In dorsal
view, regularly subelliptical; greatest breadth medial and about 45% the length.
Inner lamellae moderately broad anteriorly, narrower ventrally and posteriorly;

Pseudocypretta has a smooth shell, the inner lamella bears radial septa (in the right valve only),
the maxillule 3rd lobe has 2 Zahnborsten, the furca is reduced as in other cypridopsids. Because
the shell bears radial septa, this genus is considered to be transitional to the family Cyprettidae, in
which all known genera have such septa. On these grounds, Pseudocypretta likewise merits its own
subfamily, herein named Pseudocyprettinae, new subfamily.

Three fossil genera also are not included. They are Obliquopsis, Curvopsis and Cavernopsis,
all described by Malz (1977) who differentiates them from the living Cypridopsinae in his paper.

VOLUME 95, NUMBER 4 769

16 17

Figs. 1-18. Austrocypridopsis terrilli. 2-6, 12-18, male holotype AM P31528; 1, 7-11, female
paratype AM P31527. Internal lateral view female left valve; 2, Zenkers Organ; 3, Maxillule palp
segments; 4, Antennule; 5, Antenna; 6, Labrum; 7, Mandible endopod, with 6 and y bristles; 8,
Maxillar (P1) endopod; 9, Labium; 10, Mandible coxale, basis and endopod segments; 11, Maxillule
palp and lobes, with Zahnborsten; 12, Right maxillar palp; 13, Left maxillar palp; 14, Walking leg
(P2); 15, Cleaning limb (P3); 16, Chitinous mandibular supports with mandibular scars (note how
short these supports are); 17, Furca; 18, Hemipenis. Magnifications: Fig. 1, x 280; Figs. 2-18, x450.

770 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

line of concrescence narrow anteriorly and posteriorly, broadening medioven-
trally; radial pore canals short and straight; normal pore canals numerous, con-
centrated anteriorly and ventrally, scattered dorsally, with relatively few medially
on each valve, of simple rimmed type; muscle scars cypridopsid, comprising 4
large adductors (the posterior adductor divided), 2 large mandibulars, a small,
upper, frontal scar and several dorsal scars (the latter not illustrated); hinge a
simple ridge and groove. Spermatic vesicles of male looped posteriorly and an-
teriorly; ovigerous duct of female J-shaped, the arch of the J being posteroventral,
and extending from the posteroventral region to just behind and above the ad-
ductor scars. Eye naupliar, eyecups fused.

Antennule 7-segmented; length ratios of segments 35:20:18:11:11:9:14; dorsal/
ventral chaetotaxy of segments 1/2, 1/0, 1/0, 2/2, 2/2, 3/2, and 3 (or 4) terminal
bristles. Antennal coxa bearing 3 bristles (1 proximal and 2 ventrodistal); basis
without bristles; exopod the normal small basal plate with | long and 2 short
bristles; endopod 3-segmented; length ratios of endopod segments 12:8:nearly 3;
first endopod segment with long ventroproximal sensory bristle, 5 reduced ‘‘na-
tatory”’ bristles dorsodistally (4 equally long, one shorter), and long ventrodistal
bristle; chaetotaxy of second segment normal; 4 terminal claws, in males one of
these more slender and barbate (the normal sexual dimorphic character). Labrum
pointed anteroventrally. Detail of rake-like organs could not be observed, but
these are certainly present. Mandible coxa powerful; epipod with 5 Strahlen plus
one small bristle; chaetotaxy of basis and endopod segments includes a slender
a bristle, plump £6 bristle and long y bristle with spiky hairs. Maxillule epipod
with 15 Strahlen and 4 downwards pointing bristles; both segments of palp elon-
gate; third lobe with 2 smooth Zahnborsten. Maxilla (P1) endopod a simple lobe
with 3 terminal bristles in female, modified in male as a clasping palp, rather
similar on both sides; epipod absent. Walking leg (P2) normal; endopod segments
with rather long ventrodistal bristles, terminal claw powerful. Cleaning limb (P3)
with small terminal segment. Furca relatively powerful; proximally broad, ter-
minating in single strong claw, with small annulated bristle on lower margin of
ramus near base of claw. Chitin supports not observed. Hemipenis as illustrated.
Zenkers Organ with 14 whorls.

Dimensions
Length Height Breadth

1. Adult female 0.76 mm 0.38 mm 0.34 mm
2. Adult female 0.78 mm 0.38 mm 0.34 mm
3. Adult female 0.76 mm 0.40 mm 0.34 mm
4. Adult female 0.76 mm 0.40 mm 0.34 mm
5. Adult female 0.76 mm 0.40 mm 0.34 mm
6. Empty female LV 0.74 mm 0.38 mm —

7. Juvenile female 0.70 mm 0.36 mm 0.30 mm
8. Adult male (holotype) 0.74 mm 0.38 mm 0.32 mm
9. Juvenile male 0.66 mm 0.34 mm 0.26 mm

VOLUME 95, NUMBER 4 Wis

Acknowledgments

I am grateful to Dr. J. K. Lowry, Curator of Crustacea, The Australian Mu-
seum, for suggesting that I look at the material and providing bench space and
laboratory facilities during the May 1980 International Crustacean Congress at
the Museum. Mrs. M. Nichol typed the paper.

Literature Cited

Hartmann, G. 1964. Asiatische Ostracoden.—Internationale Revue der Gesamten Hydrobiologie,
Systematische Beihefte 3:1—155S.

Lofthouse, P. 1967. Cladocera, Ostracoda, and freshwater Copepoda.—B.A.N.Z.A.R.E. Reports,
Ser. B (Zollogy, Botany) 8(7): 141-144.

Malz, H. 1977. Cypridopsine Ostracoden aus dem Tertiar des Mainzer Beckens.—Senckenbergiana
Lethaea 58(4/5):219-261.

McKenzie, K.G. 1972. Results of the speleological survey in South Korea 1966 XXII. Subterranean
Ostracoda from South Korea.—Bulletin of the National Science Museum, Tokyo 15(1):155-
166.

Schornikov, E.I. 1980. Ostracodes in terrestrial biotopes.—Zoologicheskii Zhurnal 59(9): 1306-1319
[in Russian].

Smith, W. A., and R. L. Sayers. 1971. Entomostraca. In E. M. van Zinderen Bakker Sr., J. M.
Winterbottom, R. A. Dyer (Eds.), Marion and Prince Edward Islands pp. 361-372. A. A.
Balkema: Cape Town.

Riverina College of Advanced Education, Wagga Wagga, New South Wales,
Australia.

PROC. BIOL. SOC. WASH.
95(4), 1982, pp. 772-773

EVOPLOSOMA VIRGO, A NEW GONIASTERID STARFISH
(ECHINODERMATA: ASTEROIDEA) FROM THE
GULF OF MEXICO

Maureen E. Downey

Abstract.—Evoplosoma virgo, new species, is described from deep water in
the Gulf of Mexico. It is the second species of this rare genus known from the
Atlantic.

Among the starfishes collected during a survey of the benthic fauna of the Gulf
of Mexico by Dr. Willis E. Pequegnat, Texas A&M University, was a single
specimen of an undescribed asteroid of the genus Evoplosoma (family Gonias-
teridae). The specimen was sent to the National Museum of Natural History only
a few weeks after the description of another new species of this genus, E. scorpio
Downey, 1981, was published. Evoplosoma scorpio, from Rockall Tough and off
the mouth of the English Channel, was the first Atlantic representative of this
genus; other species of the genus are from Hawaii and the Indian Ocean.

Evoplosoma virgo, new species
Fig. |

Holotype.—USNM E24285.

Type-locality.—Alaminos Station 71A8-8, 30 July 1971, northwestern Gulf of
Mexico, 26°08’N, 92°43'W, 2056 m.

Description.—Disc large, inflated; arms 5, long, more or less square in cross-
section, narrow; abactinal plates small, discoidal, covered with uniform ensac-
culate granules; papular pores between plates single, fairly large; pedicellariae
scattered, short, rounded, barely distinguishable from granules; abactinal surface
overhanging marginals interradially; marginal plates large but quite thin, square,
conspicuous, equal, opposite, covered with granules like those of abactinal plates,
many, particularly distally, bearing a short, stout, conical spine; actinal areas
rather small, plates slightly larger than abactinal plates, with coarser granulation,
many plates bearing a large, flattened felipedal pedicellaria; row of actinal plates
adjacent to adambulacral series extending almost to end of arm; admbulacral
plates with more or less straight furrow margin bearing 4-5 crowded, flat, truncate
spines and, behind, a large, flattened felipedal pedicellaria aligned parallel to
furrow, surrounded by bare space and ring of angular granules; mouth plates
concealed by thick membrane (in holotype); madreporite small, covered with
thick membrane, midway between disc center and margin. R = 115 mm, r = 33
mm, R/r = 3.5/1; number of superomarginals = 26, R/SM = 4.4/1.

Color.—Reddish brown (dried).

Etymology.—The species is named for the constellation Virgo.

Discussion.—This species lacks the abactinal spines, spinelets, or tubercles
present in other species of Evoplosoma. The demarcation between disc and arms
is less abrupt than in E. forcipiferum Fisher, from Hawaii, and E. scorpio Down-

VOLUME 95, NUMBER 4 WD

.7/5 mm.

Fig. 1. Evoplosoma virgo: A, Two adambulacral plates; B, Two superomarginal plates.

ey, but much more so than in E. augusti Koehler, from the Indian Ocean. The
membrane covering the individual granules is apparently thicker than that of E.
scorpio, more like the thick membrane of E. forcipiferum, but as the specimen
was dried when received, it is difficult to be sure of this comparison. Evoplosoma
virgo differs from E. scorpio in having quite flattened, compressed, truncate,
adambulacral furrow spines, but they are not the peculiarly flattened, thin furrow
spines with expanded tips present in E. forcipiferum. The tiny rounded abactinal
pedicellariae and the large, rectangular, very flat actinal pedicellariae are unique
to E. virgo.

Acknowledgments

I thank Dr. Willis Pequegnat and Dr. Linda Pequegnat for sending me this
unusual specimen. Thanks are also due to Ailsa M. Clark of the British Museum
(Natural History) and David L. Pawson of the National Museum of Natural His-
tory for reading the manuscript and offering valuable suggestions.

Literature Cited

Downey, M.E. 1981. A new goniasterid seastar, Evoplosoma scorpio (Echinodermata: Asteroidea),
from the northeastern Atlantic.—Proceedings of the Biological Society of Washington 94:561-
563.

Fisher, W. K. 1906. The starfishes of the Hawaiian Islands.—Bulletin of the U.S. Fish Commission
for 1903, 23(3):987—-1130, pls. 1-49.

Koehler, R. 1909. Astéries recueillies par | Investigator dan 1’Océan Indien. I. Les Astéries de Mer
profonde. Calcutta: Indian Museum, Pp. 5—143, pls. I-XIII.

Department of Invertebrate Zoology, National Museum of Natural History,
Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
95(4), 1982, pp. 774-780

PARAPROVINCIALISM: REMNANTS OF
PALEOPROVINCIAL BOUNDARIES
IN RECENT MARINE MOLLUSCAN

PROVINCES

Edward J. Petuch

Abstract.—Paraprovincialism is a biogeographical pattern which takes into ac-
count Neogene paleoprovincial boundaries that have persisted into the Recent
and have been superimposed onto modern provincial arrangements. This pattern
is manifested as sets of mutually exclusive, congeneric species pairs that abut
along sharply defined boundary lines. An analysis of the paleontology and paleo-
zoogeography of the Caribbean region shows that these Recent species pairs are
not true siblings and that their ancestors arose in separate American paleoprov-
inces; the southern member having a Gatunian ancestor and the northern member
having a Caloosahatchian ancestor. Further analysis shows that these Pliocene
ancestral pairs were true interprovincial siblings and these are referred to as
‘‘ancestromas.’’ The paleozoogeography of the ancestromas is preserved in the
Recent by descendant species that have retained the original distributional pat-
terns. Paraprovincialism is found in other provinces, and examples are given for
the Recent central South Pacific and Indo-Malayasian regions. In areas where
the fossil record is poorly preserved, paraprovincialism may be a useful tool for
the reconstruction of Neogene provincial boundaries.

The research that led to the discovery of geographical heterochrony and relict
Pliocene molluscan faunas in the Caribbean (Petuch 198la, b, 1982) has also
brought to light a number of other enigmatic biogeographical patterns in that
region. One of the most interesting of these is a previously unstudied phenomenon
that involves a bipartite distributional pattern within many Caribbean gastropod
genera. As is most often the case, the ranges of two congeners are mutually
exclusive and will abut along a very well defined boundary line. When found in
a province, this pattern occurs in whole suites of species. |

Detailed analyses of faunal lists from other molluscan provinces, such as the
Indo-Pacific, have also revealed similar patterns for many genera. Since these
mutually exclusive species pairs occur too frequently to be simply random events
within a single province, I feel that they represent a real biogeographic phenom-
enon that is here referred to as “‘paraprovincialism.’’ By using examples of para-
provincialism in the Recent Caribbean gastropod fauna and linking these to the
fossil record, I will attempt to explain this problematical pattern of mutually
exclusive species pairs and abrupt faunal shifts.

Paraprovincialism in the Caribbean

Malacologists have generally recognized that the Caribbean Molluscan Prov-
ince can be divided into northern and southern components, each with indicator
congeners (Warmke and Abbott 1962:3—21, 319-328; Vermeij 1978:227—236). The

VOLUME 95, NUMBER 4

Table 1.—Examples of paraprovincialism in the Caribbean Province.

Restricted to northern Caribbean

Tegula lividomaculata (C. B. Adams)
Astraea americana (Gmelin)

Murex anniae McGinty

Murex cabritii Bernardi

Murex tryoni Hidalgo

Siratus articulatus Reeve
Dolicholatirus cayohuesonicus (Sowerby)
Latirus cariniferus (Lamarck)
Leucozonia ocellata Gmelin

Oliva reticularis Lamarck

Olivella mutica (Say)

Persicula fluctuata (C. B. Adams)
Prunum carneum (Storer)

Enaeta cylleniformis (Sowerby)
Conus jaspideus Gmelin

Conus magellanicus Hwass

Conus spurius atlanticus Clench

Restricted to southern Caribbean

Tegula viridula (Gmelin)

Astraea tecta (Lightfoot)

Murex olssoni E. Vokes

Murex donmoorei Bullis

Murex blakeanus E. Vokes
Siratus springeri (Bullis)
Dolicholatirus ernesti (Van Jutting)
Latirus bernadensis Bullock
Leucozonia lineata Usticke

Oliva oblonga Marrat

Olivella petiolita (Duclos)
Persicula interruptolineata (Megerle)
Prunum prunum (Gmelin)

Enaeta guildingi (Sowerby)

Conus puncticulatus Hwass

Conus beddomei Sowerby

Conus lorenzianus Dillwyn

WTS

ranges of these species pairs, some of which are listed here in Table 1, neatly
bisect the Caribbean into separate zoogeographic entities with little faunal over-
lap. Some zoogeographers, such as Briggs (1974:67—76), further divided the Ca-
ribbean into subprovinces based on this bipartite pattern. The boundaries of each
subregion were thought to have resulted from ecological differences inherent in
insular versus continental components of a single province.

All of these biogeographic schemes, regardless of their bases, delineate faunal
changes in approximately the same areas. In the Caribbean Sea, an abrupt shift
in species compositions is found along the Honduras-Nicaragua coastline in the
west and in the Lesser Antilles in the east (Fig. 1). A classic example is seen in
the Conus jaspideus-C. puncticulatus species pair (Walls 1979:817—-820). The two
Species coexist on islands from Martinique to the Grenadines, but C. jaspideus
is the only member found north of this zone and C. puncticulatus is the only one
to the south.’ The Oliva reticularis-O. oblonga species pair is also a good ex-
ample, with O. reticularis being found only north of the Leeward Islands and O.
oblonga being found only south of these islands and being most prevalent along
northern South America. Along with many other northern members of species-
pairs, Oliva reticularis and Conus jaspideus are always found together in shallow,
sandy areas of the northern Caribbean and Greater Antilles. These sympatric
Species are replaced by Oliva oblonga and Conus puncticulatus in the same
environments in the southern Caribbean.

A review of the paleontological literature of the western Atlantic (references
listed at the end of Table 2) showed that each member of a Recent Caribbean
species pair descended from a separate ancestral species and not from a single

1 The Brazilian variants of “‘C. jaspideus’’ mentioned by Walls actually are a similar, but different,
Species that is not related to this particular species-pair. ‘“‘Conus jaspideus’’ from the islands off
northern Venezuela are dwarf C. mindanus.

776 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Map of the Recent Caribbean Sea showing the approximate position of the line of abrupt
faunal shift that separates the Caribbean Province into northern and southern components.

common ancestor. Furthermore, many of these separate ancestral forms appear
to have been members of their own species pairs in the Pliocene, indicating that
the common ancestor for all must date from an earlier time. I will here refer to
these ancestral interprovincial sibling species pairs, which were also mutually
exclusive in their ranges, as “‘ancestromas.’’ Some of the Caribbean ancestromas
and their living descendant pairs are listed in Table 2. The key to interpreting
paraprovincialism, therefore, lies in detecting ancestromas in the fossil record.

For each Caribbean ancestroma that was examined, one member was found
to have been restricted to the Gatunian Molluscan Province and one to the Ca-
loosahatchian Molluscan Province (Petuch 1982) (Fig. 2). The Gatunian species
gave rise to the Recent southern Caribbean descendant while the Recent northern
Caribbean member of the pair arose from the Caloosahatchian member of the
ancestroma. The Gatunian and Caloosahatchian Provinces, together, gave rise to
the modern Caribbean fauna (Petuch 1982) and it is important to note that their
boundaries also ran somewhere along what is now the Nicaragua coastline and
the Lesser Antilles.

Paraprovincialism in the Caribbean, then, appears to represent ghosts of pre-
cursor paleoprovincial boundaries that have persisted, in secondary form, into
the Recent and have been superimposed onto modern zoogeographic patterns.
The exact mechanism for the formation of paraprovincialism, however, can only
be conjectured at this time. Most probably, the answer will be an ecological one,

VOLUME 95, NUMBER 4 UT

Table 2.—Some Recent species pairs and their Pliocene ancestromas. (N) = restricted to Recent
northern Caribbean; (S) = restricted to Recent southern Caribbean; (C) = restricted to Caloosahatch-
ian Province; (G) = restricted to Gatunian Province.

Recent pair Ancestroma
(N) Astraea americana (Gmelin) (C) Astraea precursor Dall
(S) Astraea tecta (Lightfoot) (G) Astraea aora Woodring
(N) Murex rubidum F. C. Baker (C) Murex aff. recurvirostris (n. sp.)
(S) Murex messorius Sowerby (G) Murex polynematicus Brown
(N) Latirus cariniferus (Lamarck) (C) Latirus tessellatus Dall
(S) Latirus bernadensis Bullock (G) Latirus anapetes Woodring
(N) Oliva reticularis Lamarck (C) Oliva carolinensis Conrad
(S) Oliva oblonga Marrat (G) Oliva brevispira Gabb
(N) Olivella mutica (Say) (C) Olivella dodona Olsson and Harbison
(S) Olivella petiolita (Duclos) (G) Olivella venezuelensis Weisbord
(N) Enaeta cylleniformis (Sowerby) (C) Enaeta isabellae (Maury)
(S) Enaeta guildingi (Sowerby) (G) Enaeta perturbatrix (Maury)
(N) Persicula fluctuata (C. B. Adams) (C) Persicula ovula (Conrad)
(S) Persicula interruptolineata (Megerle) (G) Persicula mareana Weisbord
(N) Prunum carneum (Storer) (C) Prunum onchidella (Dall)
(S) Prunum prunum (Gmelin) (G) Prunum macdonaldi (Olsson)
(N) Conus jaspideus Gmelin (C) Conus waccamawensis B. Smith
(S) Conus puncticulatus Hwass (G) Conus caboblanquensis Weisbord
(N) Conus spurius atlanticus Clench (C) Conus cherokus Olsson and Petit
(S) Conus lorenzianus Dillwyn (G) Conus longitudinalis Pilsbry

Ancestromas compiled from Dall 1889, 1890; S. Hoerle and E. Vokes 1978; Jung 1965; Olsson 1922;
Olsson and Harbison 1953; Olsson and Petit 1964; Pilsbry 1922; Weisbord 1962; Woodring 1928.

taking into account competitive exclusion at the provincial level. The relationship
between paleoprovinciality, ancestromas, and Recent species pairs is shown
schematically in Fig. 3.

Predictive Aspects Of Paraprovincialism

Many of the ‘‘sibling species’’ pairs in the Caribbean such as Oliva reticularis-
O. oblonga and Conus jaspideus-C. puncticulatus are actually pseudosiblings and
now can be seen to be separate offshoots of Pliocene ancestromas. For example,
these two pairs arose from the Pliocene Oliva carolinensis-O. brevispira and
Conus waccamawensis-C. caboblanquensis ancestromas, or some other closely
related species, and have retained the original distributional patterns of their
ancestral species complexes. Altogether, these two and many other Recent gas-
tropod pairs, outline the boundaries of the precursor provinces. In the case of
the Caribbean region, with a well-preserved fossil record, reliance on secondary
inferences such as paraprovincialism is not necessary for paleoprovincial recon-
structions. In areas where the fossil record is poorly preserved, however, para-
provincialism in the modern fauna may be of use to paleontologists in mapping
paleoprovincial boundaries.

Many examples of paraprovincialism are also found in the Recent Indo-Pacific

778 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Map of the Caribbean region in the Pliocene showing the distributions of the Gatunian
Province (oblique lines rising to the left) and Caloosahatchian Province (oblique lines rising to the
right) (after Petuch 1982).

Molluscan Province. The Cypraeidae are of particular interest here and are useful
indicator organisms. In the central South Pacific, a boundary line is sharply de-
lineated, in a north-south direction, from the Phoenix-Canton Atolls to east of
Samoa and west of Tonga. To the east of this line are common, wide-ranging
species such as Cypraea serrulifera, C. cumingii, and C. obvelata. To the west
of this line are the very similar C. minoridens, C. catholicorum, and C. annulus.
Along this line, no two members of a species pair have ever been found to be
sympatric (Burgess 1970).

Many other gastropods follow the same distributional patterns as those of these
cypraeids, and their combined ranges may reflect the paleoprovincial distribution
of the central South Pacific. Unfortunately, since the fossil record of this area is
so poorly known, the existence of ancestromas for most of the species pairs can
also only be conjectured at this time. The presence of paraprovincialism, how-
ever, implies that the Recent central South Pacific molluscan fauna, like that of
the Caribbean, originated from the fusion of two paleoprovinces in the Pleisto-
Cenc: |

The Malaysian Archipelago region of the Indo-Western Pacific also represents
an area of faunal shift in a paraprovincial situation. Here, the ranges of such
western (Indian Ocean) species as Strombus decorus, Strombus sibbaldi, Lambis
indomaris, Harpa crassa, Oliva ponderosa, Conus maldivus, Conus sumatrensis,
and Conus fuscatus abut along those of, respectively, the eastern (Pacific Ocean)
‘‘sibling’’ species Strombus luhuanus, Strombus plicatus, Lambis scorpio, Harpa

VOLUME 95, NUMBER 4 779

Fig. 3. Schematic diagram of the formation of paraprovincialism in the Recent Caribbean. A =
ancestral species at time T,; B,, C; = components of descendant ancestroma at time T, (Pliocene);
B,, C, = descendant species pair at time T; (Recent). Dashed line between members of ancestroma
represents paleoprovincial boundaries; solid line between Recent species pair represents sharp break
in distributions.

amouretta, Oliva miniacea, Conus generalis, Conus vexillum, and Conus imper-
ialis. These and many other Indian Ocean-Western Pacific species pairs may
reflect ancestromas that resulted from provincial differentiation during sea level
fluctuations in the Upper Pliocene and Lower Pleistocene.

Judging from the paraprovincialism of the central and western Pacific, it now
appears that the Indo-Pacific Molluscan Province, in total, may have resulted
from the post-Pleistocene combination of at least three distinct Neogene prov-
inces. As seen in this example and in the previously-mentioned Caribbean region,
the search for paraprovincialism in other zoogeographic regions may prove to be
a useful tool for the reconstruction of Neogene marine provinciality at the world-
wide level.

Acknowledgments

I thank Dr. Geerat J. Vermeij, University of Maryland, and Dr. M. G.
Harasewych, University of Delaware, for comments and critical review of
the manuscript. Special thanks go to Prof. Arthur J. Cain, University of Liver-
pool, England, for encouragement and for helping me formulate some of the ideas

780 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

on paraprovincialism. Thanks also to Mrs. Linda Flanigan for typing the manu-
script. The research for this paper was supported by National Science Foundation
postdoctoral research fellowship PDF-8166032.

Literature Cited

Briggs, John C. 1974. Marine zoogeography.—McGraw-Hill Book Co., New York. xi + 475 pp.

Burgess, C. M. 1970. The living cowries.—A. S. Barnes and Company, South Brunswick, New
York. 389 pp.

Dall, William H. 1889. Contributions to the Tertiary fauna of Florida, with especial reference to
the Miocene silex-beds of Tampa and the Pliocene beds of the Caloosahatchie River. Part 1.—
Transactions of the Wagner Free Institute of Science of Philadelphia 3: 1—200.

—. 1890. Contributions to the Tertiary fauna of Florida, with especial reference to the Miocene
silex-beds of Tampa and the Pliocene beds of the Caloosahatchie River. Part 2.—Transactions
of the Wagner Free Institute of Science of Philadelphia 3:201-473.

Hoerle, Shirley E., and Emily H. Vokes. 1978. A review of the volutid genera Lyria and Falsilyria
(Mollusca: Gastropoda) in The Tertiary of the western Atlantic.—Tulane Studies in Geology
and Paleontology 14:105—130.

Jung, Peter. 1965. Miocene mollusca from the Paraguana Peninsula, Venezuela.—Bulletins of Amer-
ican Paleontology 49(223):389-6S2.

Olsson, Axel A. 1922. The Miocene of northern Costa Rica, with notes on its general stratigraphic

relations.—Bulletins of American Paleontology 9(39): 1-168.

, and Anne Harbison. 1953. Pliocene mollusca of southern Florida, with special reference to

those from north Saint Petersburg.—The Academy of Natural Sciences of Philadelphia, Mono-

graph 8: 1-438.

, and Richard E. Petit. 1964. Some Neogene mollusca from Florida and the Carolinas.—

Bulletins of American Paleontology 47(217):509-575.

Petuch, Edward J. 198la. A relict caenogastropod fauna from northern South America.—Malaco-
logia 20(2):307-347.

—. 1981b. A volutid species radiation from northern Honduras with notes on the Honduran
Caloosahatchian secondary relict pocket.—Proceedings of the Biological Society of Washing-
ton 94:1110-1130.

—. 1982. Geographical heterochrony: Contemporaneous coexistence of Neogene and Recent
molluscan faunas in the Americas.—Palaeogeography, Palaeoclimatology, and Palaeoecology
(37):277-312.

Pilsbry, Henry A. 1922. Revision of W. M. Gabb’s Tertiary mollusca of Santo Domingo.—Pro-
ceedings of the Academy of Natural Sciences of Philadelphia 73:305—435.

Vermeij, Geerat J. 1978. Biogeography and adaptation.—Harvard University Press, Cambridge,
Massachusetts. 332 pp. .

Walls, Jerry G. 1979. Cone shells, a synopsis of the living Conidae.—Tropical Fish Hobbyist Pub-
lications, Neptune City, New Jersey. 1011 pp.

Warmke, Germaine L., and R. Tucker Abbott. 1962. Caribbean seashells.—Livingston Publishing
Company, Narberth, Pennsylvania. 348 pp.

Weisbord, Norman E. 1962. Late Cenozoic gastropods from northern Venezuela.—Bulletins of
American Paleontology 42(193): 1-672.

Woodring, Wendell P. 1928. Miocene mollusks from Bowden, Jamaica. Part 2. Gastropods and
discussion of results.—Contributions to the Geology and Palaeontology of the West Indies,
Carnegie Institution of Washington, Publication No. 385:1—564.

Department of Zoology, University of Maryland, College Park, Maryland 20742.

PROC. BIOL. SOC. WASH.
95(4), 1982, pp. 781-787

A EUNICID POLYCHAETE FROM A WHITE SMOKER
Kristian Fauchald

Abstract.—Eunice pulvinopalpata, a new species of eunicid polychaete is de-
scribed from the geothermal rift-area at 21°N off western Mexico. A brief review
of eunicids described from slope and abyssal depths is given.

During ALVIN Dive 1214 to the East Pacific Rise geothermal vent area at 21°N
in the Pacific Ocean off western Mexico four specimens of a large eunicid poly-
chaete were collected at the base of a white smoker. The material was sent to
me for identification and it proves to belong to a previously undescribed species.
This paper is OASIS Expedition Contribution #3.

Eunice pulvinopalpata, new species
Fig. 1

Material examined.—Pacific Ocean off western Mexico, 20°S5S0’N, 109°06’W,
2633 m, ALVIN dive 1214, sample #7, 20 April 1982, at base of white smoker
(Holotype, USNM 74304, | paratype USNM 74305, | paratype, British Museum
(Natural History), | paratype, Allan Hancock Foundation, Los Angeles, Poly
ISD

Description.—The holotype is an incomplete specimen with 348 setigers that
is 450 mm long and 12 mm wide including parapodia. A complete paratype (USNM
74305) has 353 setigers, but is shorter at 320 mm; the posterior end is regenerating.
All specimens are white as preserved in alcohol; eyes are absent. The body is
cylindrical; it is abruptly tapering anteriorly and slowly tapering posteriorly. The
dorsal side is arched; the ventral surface is flattened. The parapodia are attached
at the junction between the two surfaces. The pygidium of the complete paratype
(USNM 74305) is a flattened pad attached obliquely over the posterior end with
the anus opening on the sloping dorsal side. One pair of short dorsolateral and
one pair of long ventrolateral anal cirri are present.

The prostomium (Fig. la—b) is a short transverse lobe with a pair of very large,
somewhat flattened pillow-shaped palps attached along the anterior edge. The
cleft between the 2 palps is shallow. The 5 occipital antennae taper from the base.
The inner lateral and median antennae are similar in length and reach the middle
of the setiger 2. The outer lateral antennae are about half as long as the other
antennae. All antennae are irregularly articulated; the median and inner lateral
antennae have up to 19 articles of which the basal articles make up about 14 of
the total length of the antennae. The outer lateral antennae have 11 or 12 articles.

The peristomium (Fig. la) is clearly divided into 2 rings ventrally and dorsally;
laterally the separation between the 2 rings is indistinct. The total length of the
peristomium is about the same as the length of the 3 first setigers combined; the
first ring makes up about 34 of the total length of the peristomium. A pair of
peristomial cirri is attached dorsolaterally on the second ring. They reach nearly
to the tip of the palps. Each cirrus is slender and tapering and has about 12
irregular articles.

782 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

\
{ \
\s\ \
ae
VV ‘\ .
AWA atll,
WW IY aN IN! \yyt | SY, /
es 7 MeeFG aN Remedies tl 1114
AI ON SS I

d oe AUN

a 5mm | rf
Gl 0.05 mm
e 2mm

d 0.1mm

b 5mm

NSS ON e f

Fig. 1. Eunice pulvinopalpata, holotype. a, Anterior end, lateral view; b, Anterior end, dorsal
view; c, Pectinate seta, parapodium 100; d, Subacicular hook, parapodium 100; e, Parapodium 100,
anterior view; f, Compound hook, parapodium 100.

Branchiae (Fig. le) are present from setiger 3 or 4 to near the end of the
specimens. The first branchia is a short, slender filament; all other branchiae are
pectinate. The second branchia has 3 filaments; the number of filaments increases
rapidly over the next several segments so that by setiger 20 about 23 branchial

VOLUME 95, NUMBER 4 783

filaments are present. The maximum number of filaments is 25 and this number
is maintained in most setigers back to about setiger 275, after which the number
of filaments is reduced to about 15. The filaments are arranged on one side of a
distinct, rather thick branchial axis, but the branchiae are flaccid and are usually
folded down over the dorsum. Where the branchiae are best developed the pairs
meet at the dorsal midline.

All parapodia are similar. The acicular lobe (Fig. le) is distally rounded with
the acicula, usually 3 in number, emerging on the dorsal side. Pre- and postaci-
cular lobes are transverse folds covering the bases of the setae. The ventral cirri
are thick, and somewhat inflated basally, especially in median setigers, in both
anterior and posterior setigers they are digitiform. Dorsal cirri are very long,
especially in the anterior 12 of the body, where they are considerably longer than the
branchiae and reach across the body. Each cirrus is smooth, slender, and tapers
from its base.

Setae include compound hooded hooks, slender limbate setae, pectinate setae,
subacicular hooks, and acicula. The acicula are dark brown or black and taper
to bluntly pointed tips. Limbate setae are present in thick, supra-acicular fascicles
in all setigers; each seta is slender and has a very narrow limbation and a super-
ficial resemblance to a true capillary seta. Pectinate setae are present in thick
fascicles just superior to the limbate setae. Each pectinate seta (Fig. Ic) is distally
asymmetrical with a very thick marginal spine on one side. The tips of the 10-
11 distal teeth form an oblique line running from the marginal spine down toward
the other side of the seta; the bases of the teeth are all at the same level, so those
closest to the marginal spine are the longest. All pectinate setae are flat. Com-
pound hooks are present in thick fascicles inferior to the acicula. Each hook (Fig.
1f) has a short, bidentate appendage in which both teeth are of roughly the same
size; the hoods are short and blunt. The shaft is only slightly expanded near the
tip and is smooth. Subacicular hooks are first present from setiger 68 in the
holotype (from 62-72 in the paratypes). Each hook (Fig. 1d), which emerges on
the anteroventral side of the parapodium, in front of the ventral cirrus, is basally
dark brown or black, tapering to a translucent, nearly clear tip. The 2 teeth both
point distally and the proximal tooth is thicker than the distal one. The hood is
short and truncate.

The maxillary apparatus (dissected in a paratype, USNM 74305) is extremely
well sclerotinized and completely black, except for very narrow clear margins
along the cutting edges. All teeth are very small and even. The maxillary formula
is 1+1, 17+17, 18+0, 12+12 and 1+1. Each maxilla V is a sharply pointed fang.

Eunice pulvinopalpata belongs to group B2 as groups within the genus Eunice
were defined by Fauchald (1970:203,205). Other species in this group with well
developed branchiae include E. aphroditois (Pallas, 1788:229; see also Fauchald
1970:24), E. aphroditois djiboutensis Gravier (1900:224—229), E. argentinensis
(Treadwell, 1929:3), E. coccinea Grube (1878:153—-155), E. contingens (Cham-
berlin, 1919:260-263), E. flavofasciata Grube (1878: 155-156), E. flavopicta Izuka
(1912:121-123), E. franklini Monro (1924:56—58), E. guttata Baird (1869:350), E.
investigatoris Fauvel (1932:137—138), E. macrobranchia Schmarda (1861:130; see
also Augener 1918:310—312), E. multipectinata Moore (1911:248—251), E. nesiotes
(Chamberlin, 1919:253—256), E. reducta Fauchald (1970:39-43), E. torquata Qua-
trefages (1865:312; see also Grube 1870:293, and Fauvel 1923:401-402), E. scom-
brinis Quatrefages (1865:3 19-320; see also Grube 1870:296—297) and E. sebastiani

784 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Species of Eunice with black, bidentate subacicular hooks and pectinate branchiae with
10 or more branchial filaments where the branchiae are best developed. The columns are 1: Branchiae
first present from setiger #; 2: Branchiae last present (2 = branchiae missing in second half of body,
E = branchiae present to end of complete specimen); 3: Maximal number of branchial filaments; 4:
Subacicular hooks first present from setiger #; 5: Remarks.

Species 1 2 3 4 5
argentinensis 3) v 10 19
torquata 3 E 6-8 ? sensu Fauvel (1923)
torquata 3 E 14 v sensu Grube (1870)
pulvinopalpata 34 E DS 62-72
reducta 4 VY. 20-21 34-49
aphroditois djiboutensis 5) ? 20+ i hooks tridentate?
aphroditois 5—7 E 3i/ 15-54 sensu Fauchald (1970)
coccinea 5) YY. 9-10 33
flavofasciata 5 E 12-14 15
franklini 6 2 PH) 30-40
guttata 6 ? 10 ?
investigatoris 6 E 18-20 44-45
flavopicta 7 ? 19 ?
nesiotes 7) u 14 34 type examined
scombrinus i E ity! Lv sensu Grube (1870)
contingens 7 E 18-19 30 type examined
multipectinata Zi E 12 25
sebastiani 7 y 30 7-8
macrobranchia 7-8 E 15-17 zg sensu Augener (1918)

Nonato (1965: 133-139). Other taxa in this group are listed by Fauchald (1970:
205).

The taxa listed above are reviewed in Table | where the major morphological
features used to separate species in this group are indicated. The taxa are arranged
in order of the first occurrence of branchiae. Cearly E. pulvinopalpata, while
separable from all other species in the group, is not remarkably different from
them. Morphologically it most closely resembles E. reducta, from which it can
be separated on the greater numbers of setigers with branchiae and by the con-
siderably later start of the subacicular hooks. The types of both species are similar
in size and numbers of setigers. )

Etymology.—The specific name pulvinopalpata, derived from Latin, meaning
pillow-palped, refers to the shape of the palps.

Remarks.—Most species of the genus Eunice have been described from shallow
water. In addition to E. pulvinopalpata, 15 taxa have been described from slope
and abyssal depths; and of these taxa, two appear to be referable to other, pre-
viously described taxa, but the following 14 appear valid: E. arcturi (Treadwell,
1928:475), E. balfouriana (McIntosh, 1885:301—303), E. collini Augener (1906:
133-135), E. hawaiiensis Treadwell (1906:1166; see also Hartman 1966:216), E.
manihine Longbottom (1972:339-344), E. magellanica McIntosh (1885:265-267),
E. megabranchia Fauchald (1970:33—36), E. pauroneurata (Chamberlin, 1919:
249-253), E. prognatha McIntosh (1885:268—270), E. rosaurae Monro (1939:351—
352), E. segregata (Chamberlin, 1919:237—240), E. semisegregata Fauchald (1969:
8-10) and E. validobranchiata Monro (1937:288—289). Major morphological fea-
tures and basic distributional data for the types of these taxa based on information

VOLUME 95, NUMBER 4 785

Table 2.—Species of Eunice described from slope and abyssal depths. The columns are: |. Bran-
chiae first present from setiger #; 2: Branchiae last present to body-region indicated (4 = first third
of body, % = second third of body, 34 = three-quarter of the body and E = near end of body); 3:
Maximum number of branchial filaments; 4: Color of subacicular hooks (b = black, y = yellow); S:
Number of teeth in subacicular hooks; 6: Subacicular hooks first present from setiger #; 7: Condition
of tip of hoods of compound hooks (bl = blunt, p = pointed); 8: Depth of type locality; 9: Area of
origin (ECPac = East Central Pacific, SCPac = South Central Pacific, WCAtl = West Central At-
lantic, CCPac = Central Central Pacific, WInd = Western Indian, NInd = Northern Indian, NWAtI
= Northwest Atlantic); 10: Remarks.

Species 1 D) 3 4 5 6 7 8 9 10
arcturi 67 W% 14 ye Lo 2 bl 1152 NWAtI
balfouriana — — — b_ibi x v 946 SCPac
collini 9 Vy | i Loy ? 2 637 WCAtI
hawaiiensis 2 My) 30 vy oh 29 p 500 CCPac_ type examined
manihine 8 VY 3 We ell 2IEZS ok 42) WInd
magellanica 6 E 9 Ip Ip ? bie 380 Ss WA
megabranchia 3 7) 47 We ston 35 p 894 ECPac
pauroneurata 8 E 4 br bi 28 bl 718 © ECPac_ type examined
prognatha 10 2 5 be bi ? bl 764 WCAtI
pulvinopalpata 3 E DS DOI O22. ble O83 CRac
rosaurae 3 Vy | be bi 32 bl 760 WCAtI
segregata 3 VY WEN sh Lor 36 bi 970 EC@kac
semisegregata 3 7% 3438 y i Obit 51 besos eRac
validobranchiata 5 Vy 45 We i ? p 1046 #£xNiInd

in the original descriptions and on revisions of type material, are indicated in
Table 2. Seven species in this list belong to the flavus-bidentatus group, seven
belong to the fuscus-bidentatus group. Thus, two different groups have been
equally successful in adapting to life in slope and abyssal depths. A number of
these species originally described from relatively shallow slope areas (200-750
m) have poorly developed branchiae with only a few branchial filaments. Only a
single species among them, E. hawaiiensis, has more than 10 branchial filaments
where the branchiae are best developed. All species from waters deeper than
about 800 m have more than 10 branchial filaments except the abranchiate E.
balfouriana. All species with more than 20 branchial filaments are from areas of
known seafloor spreading or from other geologically extremely active areas, in-
cluding E. hawaiiensis, alluded to above. Since the species belong to two rather
different groups of the genus, it appears likely that the strong development of
branchiae in these taxa in some fashion relates to the specific environmental
conditions in the areas in which they occur.

Acknowledgments

I would like to thank Dr. J. Frederick Grassle, Woods Hole Oceanographic
Institute, for sending me the specimens on which this paper is based, and my
colleague, Dr. Meredith L. Jones for reviewing the manuscript.

Literature Cited

Augener, H. 1906. Reports on the results of dredging, under the supervision of Alexander Agassiz,
in the Gulf of Mexico and the Caribbean Sea, and on the east coast of the United States,1877

786 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

to 1880, by the U.S.S. Coast Survey Steamer Blake. Westindische Polychaeten.—Bulletin of

the Museum of Comparative Zoology, Harvard College 43:91—196, 8 plates.

. 1918. Polychaeta. Jn: W. Michaelsen (ed.)—Beitrage zur Kenntnis des Meeresfauna West-

Afrikas, 2(2):67—625, 6 plates.

Baird, W. 1869. Remarks on several genera of Annelides belonging to the group Eunicea, with a
notice of such species as are contained in the collection of the British Museum and a description
of some others hitherto undescribed.—Journal of the Linnean Society of London 10:341-361.

Chamberlin, R. V. 1919. The Annelida Polychaeta.—Memoirs of the Museum of Comparative
Zoology, Harvard College 48:1-514, 80 plates.

Fauchald, K. 1969. A revision of six species of the flavus-bidentatus group of Eunice (Eunicidae:
Polychaeta).—Smithsonian Contributions to Zoology 6:1—15, 6 figures.

—. 1970. Polychaetous annelids of the families Eunicidae, Lumbrineridae, Iphitimidae, Ara-
bellidae, Lysaretidae and Dorvilleidae from western Mexico.—Allan Hancock Monographs in
marine biology 5:1—335, 27 plates.

Fauvel, P. 1923. Polychetes errantes.—Faune de France 5:1—488, 188 figures.

1932. Annelida Polychaeta of the Indian Museum, Calcutta.x—Memoirs of the Indian Mu-
seum, Calcutta 12(1): 1-262, 9 plates, 40 figures.

Gravier, C. 1900. Contribution a ]’étude des Annelides polychetes de la mer Rouge.—Nouvelles
Archives des Museum de Science Naturelle, Paris series 4, 2(2):137—282, 6 plates.

Grube, A.-E. 1870. Bemerkungen uber Anneliden des Pariser Museums.—Archiv fur Naturge-
schichte, Berlin 36(1):281—352.

—. 1878. Annulata Semperiana. Beitrage zur Kenntnis der Anneliden-fauna der Philippinen
nach den von Herrn Prof. Semper mitgebrachten Sammlungen.—Mémoirs des Academie des
Science, St. Pétersbourg 25, LX and 300 pp., 15 plates.

Hartman, O. 1966. Polychaetous annelids of the Hawaiian Islands.—Occasional Papers of Bernice
P. Bishop Museum, Honolulu 23(11): 163-252, 2 figures.

Izuka, A. 1912. The errantiate Polychaeta of Japan.—Journal of the College of Science, Tokyo
30(2): 1-262, 24 plates.

Longbottom, M. R. 1972. Eunice manihine sp. nov. (Polychaeta, Eunicidae), a member of the
flavus-bidentate group from the western equatorial Indian Ocean.—Bulletin of the British Mu-
seum (Natural History) 21(8):339-344.

McIntosh, W. C. 1885. Report on the Annelida Polychaeta collected by H.M.S. Challenger during
the years 1873—76.—Challenger Reports 12:1-554, 82 plates.

Monro, C. C. A. 1924. On the polychaeta collected by H.M.S. Alert, 1881-1882. Fam. Polynoidae,
Sigalionidae and Eunicidae.—Journal of the Linnean Society of London 36:37—64, 24 figures.

—. 1937. Polychaeta.—The John Murray Expedition, 1933-34, Scientific Reports 4(8):243-321,

28 figures.

1939. Polychaeta of the Rosaura Expedition.—Novitates Zoologicae, London 41:345—354,

4 figures. :

Moore, J. P. 1911. The polychaetous annelids dredged by the U.S.S. Albatross off the coast of
southern California in 1904. Euphrosynidae to Goniadidae.—Proceedings of the Academy of
Natural Science, Philadelphia 63:234—318, 7 plates.

Nonato, E. 1965. Eunice sebastiani sp. nov. (Annelida, Polychaeta).—Boletim da Instituto Ocean-
ografico, Sao Paulo 14:133-139.

Pallas, P.S. 1788. Marinavaria nova et rarioraa—Nova Acta Academia Imperiale Petropolitanae 2:
229-249, | plate.

Quatrefages, A. de 1865. Histoire naturelle des Anneles marins et d’eau douce. Annelides et Gé-
phyriens.—Paris, Libraire Encyclopedie Roret 1:1—588.

Schmarda, L. K. 1861. Neue wirbellose Thiere beobachtet und gesammelt auf einer Reise um die
Erde 1853 bis 1857. 1.—Turbellarien Rotatorien und Anneliden. pt. 2:1-164, 22 plates and 120
figures. Leipzig.

Treadwell, A. L. 1906. Polychaetous annelids of the Hawaiian Islands, collected by the steamer
Albatross in 1902.—Bulletin of the United States Fisheries Commission 23:1145—-1181, 81 fig-
ures.

—. 1928. Polychaetous annelids from the Arcturus oceanographic expedition.—Zoologica, New
York 8:449-489, 3 figures.

VOLUME 95, NUMBER 4 787

—. 1929. Two new species of polychaetous annelids from the Argentine coast.—Proceedings
of the United States National Museum 75:1-5, 6 figures.

Department of Invertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington D.C. 20560.

PROC. BIOL. SOC. WASH.
95(4), 1982, pp. 788-792

CURIMATOPSIS MYERSI, A NEW CURIMATID
CHARACIFORM FISH (PISCES:
CHARACIFORMES) FROM PARAGUAY

Richard P. Vari

Abstract.—Curimatopsis myersi is described from the Rio Paraguay drainage
basin. This highly sexually dimorphic curimatid is readily recognizable by the
horizontally elongate midlateral spot on the caudal peduncle. The species is most
closely related to C. evelynae and C. crypticus from the Rio Amazonas and Rio
Orinoco systems and the Atlantic drainages of the Guianas. Curimatopsis myersi
is the first member of the genus (sensu stricto) known from a river system south
of the Rio Amazonas drainage basin.

In a recent revision, Vari (1982) reviewed the then-known species of the cur-
imatid characiform genus Curimatopsis and redefined that taxon along hypothe-
sized monophyletic lines. After that paper had gone to press, specimens of an
undescribed species assignable to that lineage were sent to the National Museum
of Natural History by Larry Naylor of the United States Peace Corps. The ma-
terial was collected as part of the Biological Inventory of Paraguay carried out
under the auspices of the Servicio Forestal Nacional de Paraguay. This new
species 1s the first member of Curimatopsis (sensu stricto) known from outside
of the Rio Amazonas and Rio Orinoco systems and the rivers of the Atlantic
drainages of the Guianas. All specimens appear to have been in breeding condi-
tion, with females bearing several dozen well-developed eggs. The type locality
is a Swampy area with turbid waters and floating plants. Counts and measure-
ments in the following description follow the methods outline in Vari (1982:2-3).
Values in square brackets are those of the holotype.

Curimatopsis myersi, new species
Figs. 1-2

Holotype.—National Museum of Natural History, USNM 233602, 39.3 mm
standard length (SL), female, collected by N. Scott and L. Fitzgerald, 31 October
1981, in a swamp 3 km northwest of Lima, San Pedro Department, Paraguay
(approx. 23°55’S, 56°29’W).

Paratypes.—39 specimens taken with the holotype, all females unless otherwise
noted: USNM 233601, 8 specimens, 2 males, 27.0-28.1 mm SL, 6 females (1
cleared and counterstained for cartilage and bone), 33.1—42.4 mm SL; Museu de
Zoologia da Universidade de S40 Paulo, MZUSP 20659, 1 specimen, 36.5 mm
SL; Museo de Historia Natural de Paraguay, 8 specimens, 2 males, 27.0—27.6
mm SL, 6 females, 34.5-42.2 mm SL; American Museum of Natural History,
AMNH 50002, | specimen, 40.7 mm SL; California Academy of Sciences, CAS
50707, | specimen, 41.5 mm SL; British Museum (Natural History), BMNH
1982.16.18:3, | specimen, 40.1 mm SL; University of Michigan, Museum of Zo-
ology, UMMZ 209809, 11 specimens, 33.9-43.5 mm SL; UMMZ 209810, 9 spec-
imens, males, 25.9-31.3 mm SL.

VOLUME 95, NUMBER 4 789

Fig. 1. Curimatopsis myersi, holotype, female, USNM 233602, 39.3 mm SL.

Diagnosis.—A robust, markedly sexually-dimorphic Curimatopsis species.
Largest known male 28 mm SL, largest female 42 mm SL. Curimatopsis myersi
can be distinguished from its congeners by the pronounced horizontal develop-
- ment of the dark patch of chromatophores on the midlateral surface of the caudal
peduncle. Curimatopsis myersi can be further distinguished from C. microlepis
in having 25 to 29 scales (versus 57 to 63) in a longitudinal series from the su-
pracleithrum to the hypural joint. Curimatopsis myersi is also separable from C.
macrolepis in having an anteriorly convex dorsal profile of the head (versus an-
teriorly straight), fusion of hypurals | and 2 (versus a separation of the elements),
and the postorbital portion of the head 0.40-0.44 of head length (HL) (versus
0.45—-0.53 of HL). The body depth of C. myersi (0.35—0.41 of SL) separates that
species from both C. crypticus (0.31—0.35) and C. evelynae (0.27-0.33).

Description.—Body moderately elongate, somewhat compressed, more so in
males. Dorsal profile of head slightly concave, particularly anteriorly. Dorsal
profile of body straight or slightly convex from rear of head to origin of rayed
dorsal fin, straight and posteroventrally slanted at base of dorsal fin, straight or
gently convex from base of last dorsal-fin ray to caudal peduncle. Dorsal body
surface with an indistinct median keel immediately anterior to rayed dorsal fin,
smoothly transversely rounded posterior to fin. Ventral body profile gently curved
from tip of lower jaw to caudal peduncle, convexity increased in gravid females.
Prepelvic region somewhat flattened transversely, but without distinct longitu-
dinal lateral keels. Ventral body surface posterior to pelvic fin insertion somewhat
flattened transversely. Greatest body depth at origin of rayed dorsal fin, depth
0.35-0.41 (0.35—0.37 in males, 0.37—0.41 in females) [0.39]; snout tip to origin of
rayed dorsal fin 0.49-0.53 [0.50]; snout tip to origin of anal fin 0.79-0.85 [0.82];
snout tip to insertion of pelvic fin 0.53—0.60 (0.53—0.58 in males, 0.56—0.60 in
females) [0.57]; snout tip to anus 0.76—-0.80 [0.78]; origin of rayed dorsal fin to
hypural joint 0.51—0.57 (0.51-0.54 in males, 0.53-0.57 in females) [0.55]. Rayed
dorsal fin pointed, anteriormost rays approximately 1.5 times length of ultimate
ray. Pectoral fin pointed; length of pectoral fin 0.17—0.21 [0.19], extending one-

790 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Curimatopsis myersi, paratype, male, USNM 233601, 27.0 mm SL.

half to three-quarters distance to vertical through insertion of pelvic fin. Pelvic
fin pointed, length of pelvic fin 0.19-0.26 (0.23—0.26 in males, 0.19—0.23 in fe-
males) [0.21], reaching three-quarters of distance to anal fin. Caudal fin form
sexually dimorphic, bifid in females; middle rays of caudal fin slightly lengthened
in males, fin trifid with upper lobe longest. Adipose fin well developed. Anal fin
emarginate, anteriormost branched rays twice length of ultimate ray. Caudal pe-
duncle depth markedly sexually dimorphic, 0.17—0.19 in males, 0.13-0.15 in fe-
males [0.15].

Head somewhat pointed, snout rounded, head length 0.24—0.33 [0.31]; jaws
equal, mouth terminal; snouth length 0.18—0.26 [0.25]; nostrils separated by a
distance less than longest dimension of anterior opening; anterior nares longitu-
dinally elongate, posterior transversely widened with anterior margin straight and
posterior convex; orbital diameter 0.30—0.38 [0.33]; length of postorbital portion
of head 0.40-0.44 [0.43]; gape width 0. 18—0.23 [0.20]; interorbital width 0.38—0.43
[0.40]. |

Scales in longitudinal series from supracleithrum to hypural joint, 25 to 29 [29];
3 to 6 lateral-line scales pored, canals in pored scales straight or that in last pored
scale somewhat curved ventrally; 2 or 3 series of scales extend beyond hypural
joint onto caudal-fin base; 10 to 12 [11] scales in a transverse series from origin
of rayed dorsal fin to midventral line.

Rayed dorsal-fin rays 11,9 [11-9]; anal-fin rays 11,7 or 8 [11,7]; pectoral-fin rays 11
to 13 [13]; pelvic-fin rays 1,7 or 8,1 [1,7,1].

Total vertebrae 27 (11 specimens), 28 (27 specimens), 29 (2 specimens).

Coloration.—Overall body coloration in preserved specimens yellowish-tan,
darker dorsally. Head dark dorsally with numerous chromatophores on upper lip,
antorbital region, lateral surface of maxilla and in a band across anterior portion
of lower lip. A patch of variously sized chromatophores on anterior portion of
inner surface of lower jaw. Scattered large chromatophores on operculum in
region of posteroventral terminus of opercular ridge. Body scales above horizon-
tal through lower margin of orbit outlined by small chromatophores; number of
chromatophores and extent of chromatophore field increasing dorsally; forming

VOLUME 95, NUMBER 4 791

a nearly solid pattern of pigmentation along the dorsal midline both anterior and
posterior to rayed dorsal fin. Middorsal pigmentation particularly pronounced
posterior to adipose dorsal fin; sometimes extending ventrally nearly to dorsal
margin of spot on caudal peduncle. A dusky band extends under scales of lateral
surface of body from slightly posterior of supracleithrum to caudal peduncle.
Band parallels dorsal body profile and gradually expands posteriorly. A promi-
nent, very dark, horizontally elongate patch of chromatophores on midlateral
surface of caudal peduncle. Caudal-peduncle spot extends onto base of middle
caudal-fin rays, particularly in males. Spot in both sexes irregularly ocellated;
surrounded by a lightly pigmented field approximately one scale wide. Fin rays
of all fins, particularly median fins, outlined by chromatophores. Caudal fin rosy
colored in recently preserved specimens.

Remarks.—A number of earlier researchers incorporated all curimatid species
with incompletely pored lateral lines into Curimatopsis, but authors commencing
with Myers (in Eigenmann and Myers 1929) have questioned the utility of that
character in delimiting natural lineages among small characiforms. As discussed
by Vari (1982:3-12), the union of curimatid species with an incomplete latero-
sensory canal system on the body into a single taxon actually defined a non-
monophyletic assemblage, subunits of which were more closely related to other
members of the family than to some of the other nominal Curimatopsis species.
Under Vari’s more restricted definition of Curimatopsis, the genus was consid-
ered to consist of only four species, none of which was known to occur south of

the Amazon River drainage basin. Earlier researchers had described various nom-
inal *“Curimatopsis’’ species from the La Plata drainage system or cited the pres-
ence of Curimatopsis macrolepis in the region. Meinken (1933:71) described a
species he termed Curimatopsis saladensis from the Rio Salado of Argentina.
That species has been subsequently reported from Rio Grande do Sul, Brazil
(Grosser and Hahn 1981:55) and intervening areas. ‘“‘Curimatopsis’’ saladensis
and “‘C.’’ maculatus described by Ahl (1934:240), supposedly from Argentina,
are more closely related to curimatid species outside of Curimatopsis (sensu
stricto) and do not extend the range of that genus into the La Plata drainage
system (see Vari 1982:11). Schindler (1939:275) followed by Ringuelet (1975:72)
reported Curimatopsis macrolepis from Nueva Germania, Paraguay. The data in
Schindler’s publication and the accompanying photo indicate that the specimen
reported on was neither C. macrolepis nor C. myersi, but rather a juvenile of
some non-Curimatopsis curimatid.

Relationships.—Curimatopsis myersi shares with the other four species of the
genus (sensu stricto) all eight of the shared derived characters of the caudal fin
rays, caudal osteology, suspensorium, antorbital, and dermosphenotic that char-
acterize the genus (see characters | to 8, Vari 1982:10). Within Curimatopsis, the
fusion of hypurals | and 2 into a single plate autogenous with respect to the fused
PU,+U,, the posterior contact and partial interdigitation of the fused hypural 1
and 2 with hypural 3 in males, and the pronounced vertical expansion of the
penultimate principal caudal ray of the lower lobe of the caudal fin of males are
synapomorphies linking C. myersi to C. evelynae and C. crypticus (see characters
9 to 11, Vari 1982: 10).

Etymology.—Named in honor of Dr. George S. Myers who over the course of
many years has contributed greatly to our knowledge of Neotropical characi-
forms.

792 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Acknowledgments

The specimens that served as the basis for this paper were collected during the
Biological Inventory of Paraguay carried out under the auspices of the Servicio
Forestal Nacional de Paraguay. The success of the survey was made possible by
the efforts of numerous participants from both Paraguay and the United States.
I particularly thank Larry Naylor for all his assistance in making available the
specimens which served as the basis of this paper. Reeve M. Bailey and Gerald
R. Smith (UMMZ) kindly brought additional specimens of this species to my
attention. Technical assistance in the study was provided by Robert H. Kanazawa
and Susan L. Jewett. This paper benefitted from the comments and criticisms of
Stanley H. Weitzman and Mercedes S. Foster.

Literature Cited

Ahl, E. 1934. Beschreibungen zweier neuer SUsswasserfische aus Sudamerika.—Sitzungsberichte
der Gesellschaft Naturforschende Freunde 24:238—241.

Eigenmann, C. H., and G. S. Myers. 1929. The American Characidae. 5.—Memoirs of the Museum
of Comparative Zoology 43(5):429-S15.

Grosser, K. M., and S. D. Hahn. 1981. Ictiofauna da Lagoa Negra, Parque Estadual de Itapua,
Municipio de Viamao, Rio Grande do Sul, Brazil.—tlheringia, Série Zoologia 59:45—64.
Meinken, H. 1933. Ueber einige in Letzter Zeit Eigenfiihrte Fische I1.—Blatter Fir Aquarien und

Terrarienkund 44:71-73.

Ringuelet, R. A. 1975. Zoogeografia y ecologia de los peces de aguas continentales de la Argentina
y consideraciones sobre las areas ictidlogicas de América del Sur.—Ecosur 2(3):1—122.
Schindler, O. 1939. Ueber die Fischausbeute der 3 Sudamerika-Expedition Prof. Kriegs.—Sitzungs-

berichte der Gesellschaft Naturforschende Freunde 15:268-303.
Vari, R. P. 1982. Systematics of the Neotropical Characoid genus Curimatopsis (Pisces Chara-
coidei).—Smithsonian Contributions to Zoology, no. 373, 28 pp.

Department of Vertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
95(4), 1982, pp. 793-806

ALTERNOCHELATA LIZARDENSIS, A NEW SPECIES
OF MYODOCOPINE OSTRACODE FROM THE GREAT
BARRIER REEF OF AUSTRALIA (RUTIDERMATIDAE)

Louis S. Kornicker

Abstract.—Alternochelata lizardensis is described and illustrated. The speci-
mens were collected from the Lizard Island Group, Great Barrier Reef, Australia.
The genus was known previously only from the North Atlantic.

The only previous record of a member of the Rutidermatidae Brady and Nor-
man 1896, in Australia is a listing of Rutiderma sp. by Poore et al. (1975:31, 60,
61) among the fauna of Port Phillip Bay, Victoria. According to those authors
Rutiderma sp. was collected in bottom samples at depths of 20-24 m; bottom
salinity was 34.95%co (Poore and Rainer 1975:373). Alternochelata lizardensis, the
new species described here from the Lizard Island Group of the Great Barrier
Reef, is a member of a genus recorded previously only from the North Atlantic,
viz. the Great Bahama Bank (Kornicker 1958:237), and off Mauritania (Kornicker

and Caraion 1978:66).

Alternochelata Kornicker, 1958

Type-species.—Rutiderma (Alternochelata) polychelata Kornicker, 1958, by
monotypy.

Alternochelata lizardensis, new species
Figs. 1-6

Etymology.—The specific name refers to the type-locality, Lizard Island, Aus-
tralia.

Material.—Lizard Island main lagoon; Sept—Oct 1977; undisturbed sand flat,
depth 6 m; 15 replicate small cores (each 0.0044 sq. m); cores sieved through 186
jzm mesh; specimens from all cores consolidated: | adult female, USNM 158609,
holotype; | adult male, USNM 158489, paratype. Palfrey Island; Sept-Oct 1977,
transect across sand flat from shore to reef; stations 1-3 fairly close together and
above mean datum; stations 4-8 extending 150 m across sand flat; station 8 at far
edge of flat adjacent to a coral head at edge of coral reef; stations 4-8 exposed
probably only at extreme low tides, twice a year. Sand fine to medium fine, with
small ripple marks, but not exposed to much wave action; tube-dwelling poly-
chaetes and other more-or-less sedentary animals fairly abundant; 5 large cores
(each 0.0176 sq. m) taken from each of 8 stations (ostracodes present in only 5
outer stations); cores sieved through 500 um mesh; specimens from each station
consolidated: Station 5, 1 A-1 male, USNM 158029, paratype; Station 6: 1 adult
male, USNM 158487, paratype; 3 adult females, USNM 158606A, C, D, para-
types; | ovigerous female, USNM 158606B, paratype; 5 juveniles, USNM 158606E,
paratypes; Station 7, 3 adult males, USNM 158605, paratypes; Station 8: 1 oviger-

794 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Alternochelata lizardensis, A-1 male, paratype, USNM 158029, lateral view of complete
specimen, length 1.29 mm.

ous female, USNM 158488, paratype; 7 paratypes including adult males, females,
and juveniles, deposited in The Australian Museum, Sydney. Palfrey Island; Sept—
Oct 1977, active sand cones; on sand flat at depth of 12.3 m; 10 replicate small
cores (each 0.0044 sq. m); cores sieved through 186 ~m mesh; specimens from
all replicate cores consolidated: 3 juveniles, USNM 158607, paratypes.

Description of A-I male (Figs. 1—3e).—Carapace ovoid with narrow but dis-
tinctly projecting caudal process; incisur well developed with slightly overhanging
rostrum (Figs. 1, 2a).

Ornamentation (Figs. 1, 2d): Surface with numerous shallow fossae; lateral ribs
absent; anterior margin of rostrum, and also anteroventral and ventral margins
of valve with minute scallops; scallops absent in vicinity of incisur, and also along
posterior one-fourth of ventral margin of valve, in vicinity of caudal process, and
along posterior and dorsal margins of valve; bristles numerous along edge of
rostrum, and also along anteroventral and ventral margins of valve; bristles sparsely
distributed over lateral surface; most long bristles with broad basal part.

Infold (Fig. 2a,c): Rostral infold with 8 or 9 long spinous bristles forming row,
and 1 shorter spinous bristle near inner end of incisur (Fig. 2a); 1 small bristle at
inner end of incisur not on infold (Fig. 2a); anteroventral infold with | short bristle
(just ventral to inner end of incisur) followed by space and then 6-8 longer bristles
forming row parallel to valve edge; about 7 parallel ridges present on anteroven-
tral infold dorsal to bristles; narrow list present having anterior end located on
ventral infold at point about one-third length of valve measured from posterior
end of caudal process; list continuing onto infold of caudal process and ending
on posterior infold dorsal to caudal process (Fig. 2c); 19-23 minute bristles (some
forming pairs) present on infold just outside list (anterior segment of list and some
anterior bristles not shown on Fig. 2c); infold of caudal process with small bristle
near middle (Fig. 2c).

Selvage (Figs. 1, 2b, c): Wide lamellar prolongation along anterior and ventral

VOLUME 95, NUMBER 4 795

A

rT — )

sm SS

geceveveenes0010020)
a

Fig. 2. Alternochelata lizardensis, A-| male, paratype, USNM 158029: a, Rostrum and incisur of
right valve, inside view; b, Incisur of right valve showing selvage bearing striations and marginal
fringe; c, Posterodorsal corner of right valve showing caudal process, infold, and smooth narrow
selvage divided at corner by small bristle; d, Detail of right valve from inside showing some central
adductor muscle attachment scars (lined) and fossae (unlined) (not all scars could be distinguished
from fossae with certainty); e, Left Ist antenna, medial view; f, Distal part of protopodite, endopodite,
and Ist joint of exopodite of right 2nd antenna, medial view; g, Right mandible, medial view; h, Left
maxilla, lateral view; i, 7th limb (tips of some bristles broken); j, Detail of tip of 7th limb shown in
1; k, Left lamella of furca, lateral view; 1, Anterior of body from left showing left lateral eye, medial
eye and Bellonci organ, and upper lip with single anterior process; m, Right Y-sclerite, anterior
towards right.

796 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

margins becoming narrower in vicinity of caudal process and along posterior
margin of valve; prolongation divided at inner end of incisur (Fig. 1, 2b), and
divided by minute bristle at edge of caudal process (Fig. 2c); margin of prolon-
gation with fringe of long and short hairs along anterior and ventral margins of
valve, bare in vicinity of caudal process and along posterior margin (Fig. 2c);
fringe on selvage in vicinity of incisur and along anterior margin of rostrum not
shown in Fig. 1}.

Central adductor muscle attachments (Fig. 2d): Consisting of about 18 to 24
oval scars.

Size: USNM 158029, length 1.29 mm, height 0.87 mm.

First antenna (Fig. 2e): Ist joint with slender spines on medial surface. 2nd
joint with slender spines on medial surface and dorsal margin, and with 2 bristles
(1 dorsal, | lateral). 3rd joint short, with few medial spines and 2 bristles (1 dorsal,
1 ventral). 4th joint separated from 3rd joint by distinct suture, with 4 bristles (1
dorsal, 3 ventral). Sensory bristle of Sth joint with about 6 short proximal fila-
ments, | minute filament just distal to middle, and bifurcate tip (ail filaments
drawn out to fine point). 6th joint minute, fused to Sth joint, with short, bare,
medial bristle. 7th joint: a-bristle bare, about same length as bristle of 6th joint;
b-bristle about twice length of a-bristle, with 1 small proximal filament, | minute
filament near middle, and tip with minute process; c-bristle about same length as
sensory bristle of 5th joint, with 3 small proximal filaments, 1 minute filament
distal to middle, and bifurcate tip (filaments drawn out to fine tip). 8th joint: d-
and e-bristles slightly shorter than c-bristle, bare with blunt tips; f-bristle about
one-third longer than b-bristle, with 2 short proximal filaments and bifurcate tip;
g-bristle about same length as c-bristle, with 2 short proximal filaments, | minute
filament distal to middle, and bifurcate tip (filaments drawn out to fine point).
Each branch of bifurcate terminal ends of sensory bristle, as well as c-, f-, and
g-bristles, with minute process at tip.

Second antenna (Fig. 2f): Protopodite bare. Endopodite 3-jointed: Ist joint
short with 5 short bristles (4 proximal, | distal); 2nd joint elongate with 3 or 4
ventral bristles (1 long spinous, 2 or 3 short bare); 3rd joint elongate with 3 bristles
(1 dorsal, 2 terminal). Exopodite: Ist joint elongate with minute, terminal, medial
bristle with tubular tip; bristles of joints 2-8 short, with stout spines forming
groups along ventral margins (these spines smaller in distal groups); 9th joint with
4 very short terminal bristles (ventral 2 of these with ventral spines similar to
those on joints 2-8; remaining 2 bristles bare), and 2 minute medial bristles; joints
2-8 with small slender spines along distal margins; basal spines absent.

Mandible (Fig. 2g): Coxale endite stout, bifurcate, with proximal hairs and
distal spines. Basale: medial side with 5 proximal bristles near ventral margin: 2
pectinate, 3 ringed (1 pectinate and | ringed bristle broken off on illustrated limb);
dorsal margin with 3 bristles (1 near middle, 2 subterminal); lateral side with short
bristle near middle of ventral margin. Exopodite absent. Ist endopodial joint with
medial spines and 3 ventral bristles (2 short bare, | long spinous). 2nd endopodial
joint: medial surface with spines forming rows near dorsal margin; dorsal margin
with 7 bristles; ventral margin with 2 small terminal bristles; distal end near
ventral margin with stout claw with large teeth along inner margin, and with
prolonged tip having minute teeth along inner margin; a small spade-shaped pro-

VOLUME 95, NUMBER 4 797

cess with spine at tip present medial to stout claw. 3rd endopodial joint: distal
margin with stout claw having 2 rows of teeth along inner margin; a second
narrower claw present ventral to stout claw (this claw also with teeth along inner
margin); medial surface with 3 short ringed bristles; lateral surface with 2 small
ringed bristles.

Maxilla (Fig. 2h): Endite I with 3 spinous bristles and 2 pectinate claws; endite
II with 2 bristles and 2 pectinate claws; endite III with 3 terminal pectinate claws,
3 terminal bristles, and | proximal bristle. Dorsal margins of precoxale, coxale,
and proximal part of basale, with fringe of hairs. Coxale with short, terminal,
dorsal bristle. Basale with | ventral bristle, and 2 medial bristles (1 of these near
dorsal margin, other distal and could be interpreted to be on Ist endopodial joint).
Exopodite small with 3 bristles (2 short, 1 long). Ist endopodial joint with dorsal
spines, | spinous alpha-bristle and | spinous beta-bristle. 2nd endopodial joint
with 2 stout pectinate claws and 5 slender bristles, some with marginal spines.

Fifth limb (Fig. 3a—d): Epipodial appendage with 48 bristles. Endite I with 1 or
2 short, ringed, spinous bristles and | minute unringed bristle; endite II with 3
short, ringed, spinous bristles and none or | minute unringed bristle; endite III
with 8 bristles (1 inner anterior bristle with long proximal hairs and short distal
spines adjacent to | minute unringed bristle; 2 short, slender pectinate bristles;
and 4 ringed, slender, spinous bristles). Ist exopodial joint with proximal bristle
adjacent to 2 small teeth followed by 2 curved prongs (proximal of these with
distal marginal tooth; other with 2 proximal and 3 distal marginal teeth (Fig. 3d)).
2nd exopodial joint folded over on both limbs examined; consisting of large tri-
angular tooth with 2 large teeth (distal of these with marginal tooth) along inner
margin (Fig. 3d); joint with usual posterior bristles (Fig. 3b); outer corner of
triangular tooth with small bristle with long hairs (Fig. 3a); 3rd exopodial joint
with 2 bristles (with minute marginal teeth) on inner lobe, and 2 bristles (with
long marginal hairs) on outer lobe. 4th and 5th joints fused, with total of 4 bristles.

Sixth limb (Fig. 3e): Endites I and II each with 3 bristles; endite III with 4
bristles; endite IV with 3 bristles. End joint with 8 or 9 bristles (anterior 6 or 7
bristles with either short marginal spines, or long proximal hairs and short distal
spines; posterior 2 bristles with long marginal hairs); ventral margin of end joint
linear. Two or 3 bristles with long marginal hairs present in place of epipodial
appendage. Limb without long hairs on endites or end joint.

Seventh limb (Fig. 21,j): 3 or 4 tapering bristles in proximal group (each bristle
with marginal spines and up to 3 bells); 4 tapering bristles in terminal group (each
bristle with marginal spines and up to 5 bells). Terminus with comb of 9 teeth
opposite 2 recurved teeth; comb consisting of 3 short teeth on each side of 3
slightly longer teeth; short teeth with alar projections on each side and tooth on
each side of base.

Furca (Fig. 2k): Each lamella with 9 claws; claws 1, 2, 4, 5 stout primary claws;
claws 3, 6-9, small secondary claws; claws | and 2 with long teeth along posterior
margins (small slender teeth also present); some secondary claws with minute
spines along posterior margins (not shown on illustrated limb).

Bellonci organ (Fig. 21): Elongate, broadening in middle, with rounded tip.

Eyes (Fig. 21): Lateral eye very faint, light amber color. Medial eye reddish
in preserved specimen.

798 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

eS 2

R
s
=
s
s
‘Ss
R\
2 \
AX \

Fig. 3. Alternochelata lizardensis, a—e, A-1 male, paratype, USNM 158029: a, Right 5th limb,
twisted but anterior view; b, Left 5th limb, twisted but mostly posterior view; c, Detail of tip of large
tooth of 2nd exopodial joint; d, Detail of teeth of exopodial joints 1 and 2, from b; e, 6th limb. f-l,
Adult female, paratype, USNM 158488: f, Lateral view of complete specimen showing position of 4
of 6 eggs and central adductor muscle attachments, carapace length 1.43 mm; g, Endopodite of left
2nd antenna, medial view; h, Distal part of protopodite, endopodite, and Ist joint of exopodite of
right 2nd antenna, medial view; i, Right mandible, medial view; j, Distal end of Sth joint showing
both lobes of 3rd joint (outer lobe to right) and fused 4th and Sth joints, from Figure 4d; k, Left
lamella of furca and left Y-sclerite, lateral view; 1, Anterior of body from left showing left lateral eye,
medial eye and Bellonci organ, anterior process, and upper lip with single anterior process.

VOLUME 95, NUMBER 4 729

Upper lip (Fig. 21): Lip with single anterior process.

Y-sclerite (Fig. 2m): Posterior end forming right angle; middle segment linear,
fairly long; wide angle between dorsal and ventral branches.

Description of adult female (Figs. 3f-1, 4a—e).—Similar in shape to that of a
A-1 male except caudal process not so well defined (Fig. 3f).

Ornamentation: Similar to that of A-1 male.

Infold: Rostral infold with 11 or 12 long spinous bristles forming row and 2
shorter bristles near inner end of incisur; anteroventral infold with 1 short bristle
just ventral to inner end of incisur followed by space and then 9 or 10 longer
bristles forming row; anteroventral infold also with about 6 ridges between bristles
and inner margin of infold; narrow list on infold of caudal process and posterior
end of ventral margin with 20-23 minute bristles, some in groups of 2 or rarely
3 bristles but most single; 2 or 3 small bristles on infold of caudal process between
list and posterior edge of valve.

Selvage: Similar to that of A-1 male.

Size: USNM 158488, length 1.43 mm, height 0.98 mm. USNM 158606A—D
(4 specimens): length 1.37 mm, height 0.91 mm; length 1.41 mm, height 0.94 mm;
length 1.37 mm, height 0.94 mm; USNM 158609, length 1.34 mm, height 0.91
mm.

First antenna: 4th joint with 2 long bristles on ventral margin, joints 1-4 oth-
erwise similar to those of A-1 male. 5th joint bent ventrally as on A-1 male; sensory
_bristle with 3 short proximal filaments, 1 minute distal filament and bifurcate tip
(each branch with minute terminal process). 6th joint fused to Sth joint, with
short, bare, medial bristle. 7th joint: a-bristle bare, extends slightly past tip of
bristle of 6th joint; b-bristle about twice length of a-bristle, with 1 small filament
just distal to middle, and tip with minute process; c-bristle about as long as
sensory bristle of 5th joint, with | minute proximal filament, 2 minute filaments
distal 2 middle, and bifurcate tip (each branch with minute terminal process). 8th
joint: d- and e-bristles slightly shorter than c-bristle, bare with blunt tips; f-bristle
about one-third longer than b-bristle, with 1 minute proximal filament and bifur-
cate tip (each branch with minute terminal process); g-bristle as long as c-bristle,
with 2 minute proximal filaments, 1 minute distal filament, and bifurcate tip (each
branch with minute terminal process).

Second antenna: Protopodite bare. Endopodite either 2 or 3 jointed (Fig. 3g,
h): Ist joint short, with 6 small bristles (S proximal, | distal); 2nd joint elongate
with | long spinous ventral bristle and | shorter, bare, terminal bristle (the latter
bristle on 3rd joint when dividing suture present between ventral and terminal
bristle; see (Fig. 3h)). Exopodite: Ist joint elongate, with 1 small, tubular, medial
bristle on distal margin (Fig. 3h); joints 2—9 decrease in size gradually; bristles
on joints 2-5 fairly short, with closely spaced, stout, short, ventral spines (forming
cornlike row); ventral spines on bristles of females similar to those of A-1 male
but tending to have more rounded tips; bristle on 4th joint of right limb slightly
longer than bristle of Sth joint and with short slender ventral spines near tip;
bristles of joints 6-8 of USNM 158488 and 158606B broken off near base (re-
maining part bare); 9th joint with 5 bristles (3 stout, broken off near base of
USNM 158488, 2 very small, bare); joints 2-8 with small spines forming row
along distal margin; basal spines absent. Long exopodial bristles of joints 6-9 on
USNM 158609, 158606A, C, D, unbroken, with natatory hairs, no spines.

800 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Mandible (Fig. 31): Coxale endite stout, bifurcate, with proximal hairs and distal
spines; medial hairs in row near base. Basale: medial side with 5 proximal bristles
near ventral margin (2 short pectinate, | short bare, | longer spinous and | short
spinous bristle nearer to middle of ventral margin); dorsal margin with 3 bristles
(1 near middle, 2 subterminal); ventral margin with 1 long distal spinous bristle
(base may be on lateral surface). Exopodite absent. Ist endopodial joint with
medial spines and hairs forming rows; lateral side with hairs forming single distal
row near dorsal margin; small ventral margin with | long and 2 short bristles. 2nd
and 3rd endopodial joints similar to those of A-1 male.

Maxilla: Similar to that of A-1 male.

Fifth limb (Fig. 3}, 4a—d): Endite I with 2 bristles; endite II with 3 ringed spinous
bristles and | unringed bristle; endite HI with 8 bristles (similar to those of A-1
male). Ist exopodial joint with a spinous bristle proximal to 2 stout teeth, and an
elongate pronglike tooth with 3 marginal teeth (Fig. 4c); 2 short anterior teeth at
base of pronglike tooth (Fig. 4c); 2 bristles on anterior margin of joint (Fig. 4b,
c). 2nd exopodial joint consisting of large triangular tooth with 2 large teeth along
inner margin (Fig. 4b, e); 2 long bristles on inner margin proximal to proximal
tooth (Fig. 4b, e); posterior side with | small bristle in proximal outer corner,
and 3 distal bristles (Fig. 4d, e). 3rd exopodial joint with 2 bristles (with minute
marginal teeth) on inner lobe, and 2 bristles (with long marginal hairs) on outer
lobe; 4th and Sth joints fused, with total of 5 bristles (Fig. 3)).

Sixth limb: Endite I and II each with 3 bristles; endite III with 4 bristles; endite
IV with 3 bristles. End joint with 8 ventral bristles (anterior 6 bristles with few
to many long hairs near middle and small distal spines; posterior 2 bristles with
long marginal hairs); ventral margin of end joint linear; 2 hirsute bristles in place
of epipodial appendage; trunk with few hairs along posterior edge proximal to
epipodial bristles. Limb similar to that of A-1 male (see Fig. 3e).

Seventh limb: Each limb with 6 bristles in terminal group (3 on each side) and
4 in proximal group (2 on each side; the bristles close together on comb side, and
farther apart on opposite side); each bristle with marginal spines and 4—6 distal
bells. Terminus consisting of comb with 9 mostly alate teeth opposite 3 spinous
recurved teeth.

Furca (Fig. 3k): Each lamella with 8 to 10 (usually 9) claws; claws 1, 2, 4, 5
stout primary claws; claws 3, 6-8, 9, 10 small secondary claws; claws | and 2
with stout posterior teeth; secondary claws with slender posterior spines; tips of
main claws rounded (appear worn down); claw | of right lamella anterior to claw
| of left lamella.

Bellonci organ (Fig. 31): Elongate; middle with short, broad segment delineated
by faint sutures or folds; distal part cylindrical with rounded tip.

Eyes (Fig. 31): Lateral eye small, unpigmented, with 4 or 5 minute ommatidia.
Medial eye bare, well developed, pigmented black in central portion and having
reddish tint in marginal portions when viewed with transmitted light.

Upper lip (Fig. 31): Lip with single stout anterior process.

Posterior of body: With few hairs (Fig. 3k); forming almost right angle at dorsal
end of girdle. (Girdle is posterior sclerite with ventral ends connected to posterior
ends of left and right Y-sclerites.)

Y-sclerite (Fig. 3k): Posterior end bent ventrally; middle segment linear; gai
branch obscure on specimens examined.

VOLUME 95, NUMBER 4 801

Fig. 4. Alternochelata lizardensis, a—e, adult female, paratype, USNM 158488, Sth limb: a, Right
limb, anterior view; b, Detail of 1st and 2nd exopodial joints, from a; c, Detail of Ist exopodial joint,
from b; d, Left limb, posterior view; e, Detail of 2nd exopodial joint, from d. f-i, adult male, paratype,
USNM 158487: f, Complete specimen showing right lateral eye, and central adductor muscle attach-
ments, carapace length 1.36 mm; g, Right Ist antenna (long filaments of sensory bristle of Sth joint
not shown), lateral view; h, Right lamella of furca, lateral view; i, Anterior of body from right side
showing right lateral eye (ommatidia not drawn), medial eye and Bellonci organ, and upper lip (anterior
process of upper lip bent backward on illustrated lip).

Eggs (Fig. 3f): USNM 158488 with 6 eggs in marsupium (not all shown in
illustration); USNM 158606B with 4 eggs.

Remarks: On the two ovigerous females, USNM 158488, 158606B, the long
bristles on joints 6—9 of the exopodite of the 2nd antenna are broken off; whereas,
these bristles on the four adult females, USNM 158609, 158606A, C, D, which

802 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

did not have extruded eggs, were unbroken and bore natatory hairs. The phe-
nomenon of females breaking off exopodial bristles after mating, thus rendering
them incapable of swimming, is common among the Philomedidae, but has been
previously reported in the Rutidermatidae on only one species, Alternochelata
nealei Kornicker and Caraion, 1978:73.

Description of adult male (Figs. 4f-1, -6)—Carapace more elongate than that
of adult female and with broader rostrum (Fig. 4f).

Ornamentation: Similar to that of adult female.

Infold: Not examined.

Selvage: In vicinity of rostrum and along ventral margin similar to that of adult
female, not examined elsewhere.

Central adductor muscle attachments (Fig. 4f): Similar in number to those of
A-1! male.

Size: USNM 158487, length 1.36 mm, height 0.80 mm; USNM 158489, length
1.37 mm, height 0.79 mm; USNM 158605 (3 specimens): length 1.36 mm, height
0.79 mm; length 1.38 mm, height 0.81 mm; length 1.39 mm, height 0.83 mm.

First antenna (Fig. 4g): Lateral hairs forming row on soft integument between
Ist and 2nd joints. Joints 2-4, and 6 with hairs and spines in rows. 2nd joint with
1 dorsal bristle, and 1 lateral bristle near ventral margin. 3rd joint short, with 1
dorsal bristle, and 1 ventral bristle with base on medial side of joint. 4th joint
with | spinous dorsal bristle, and 4 ventral bristles with bases on medial side of
joint. 5th joint minute, fused to 4th; sensory bristle with broad proximal part
bearing numerous long filaments (filaments not shown on illustrated limb); distal
stem of sensory bristle with 1 minute filament near middle and bifurcate tip (each
branch with minute terminal process). 6th joint about same length as 4th joint,
with | medial bristle near dorsal margin. 7th joint: a-bristle spinous, about same
length as bristle of 6th joint; b-bristle about twice as long as a-bristle, with 3
marginal filaments and tip with minute process; c-bristle very long, with 13 mar-
ginal filaments and tip with minute terminal process. 8th joint: d- and e-bristles
bare with blunt tips, longer than b-bristle; d-bristle slightly stouter and longer
than e-bristle; f-bristle very long, similar to c-bristle; g-bristle longer than d-
bristle, with 3 marginal filaments and bifurcate tip (each branch with minute
terminal process). Filaments on sensory bristle and bristles of joints 7 and 8 with
1 or 2 minute processes at tip.

Second antenna: Prodopodite bare. Endopodite 3-jointed (Fig. 5a): Ist joint
short with 6 bristles (5 proximal, | distal); 2nd joint elongate with 2 short ventral
bristles; 3rd joint elongate, reflexed on 2nd, with | proximal bristle with pointed
tip, 2 short distal bristles, and tip with few ridges. Exopodite (Fig. 5b): Ist joint
elongate, spinous, with small, medial, tubular bristle on distal margin; 2nd joint
short; 3rd joint about twice length of 2nd joint; remaining joints small, decreasing
in size distally; bristle of 2nd joint short, reaching 7th exopodial joint, with blunt
spines along ventral margin; bristles of joints 3-8 long, with natatory hairs, no
spines; 9th joint with 6 bristles (3 long and 1 medium with natatory hairs, 2 minute,
bare); joints 3-8 with slender, pointed, basal spines; 9th joint without lateral
spine; joints 2-8 with slender spines along distal margin.

Mandible (Fig. Sc): Coxale endite consisting of 2 minute spines. Basale: medial
surface spinous, with 5 proximal bristles near ventral margin and | additional
bristle near middle of ventral margin; dorsal margin with 3 bristles (1 distal to

VOLUME 95, NUMBER 4

803

Wow ie

a

My, Lp >>
Y

O008TII py,

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asnuteel

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Ay

Fig. 5. Alternochelata lizardensis, adult male, paratype, USNM 158487: a, Endopodite of left 2nd
antenna, medial view; b, Exopodite of left 2nd antenna (bristles of joints 2-8 not shown), lateral view;
c, Right mandible, medial view; d, Distal part of basale, Ist endopodial joint, and proximal part of
2nd endopodial joint of left mandible, lateral view; e, Lateral view of distal part of left mandible
showing bristles of distal ventral corner of 2nd endopodial joint, and claw and lateral bristle of end

joint; f, Right maxilla, lateral view; g, Detail from f showing exopodite and endopodite; h, 6th limb;
1, 7th limb.

middle, 2 subterminal). Exopodite absent. Ist endopodial joint with medial spines,
3 ventral bristles, and bulging process along dorsal margin (the latter process
could, with difficulty, be interpreted as being homologous to exopodite but, in
my opinion, is not). 2nd endopodial joint: medial side, ventral and dorsal margins
with spines in rows; ventral margin with bristles forming 2 distal groups, 2 bristles

804 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

\ :
ONES f a
Sean A Magar 3
a Sccayyaatte' [E 44 al SP
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Fig. 6. Alternochelata lizardensis, adult male: a-d, USNM 158487, Sth limb: a, Distal part of limb
(not all exopodial bristles shown); b, Ist exopodial joint; c, 2nd exopodial joint; d, Detail from a
showing 3rd exopodial joint (outer lobe to left), and fused exopodial joints 4 and 5. e, USNM 158489,
anterior of body from right side showing right lateral eye (ommatidia not drawn), medial eye and
Bellonci organ, and upper lip with minute process.

in proximal group, 4 in distal group (the 2 distal bristles in latter group slender
with bases on lateral side) (Fig. Se); dorsal margin with 7 bristles. End joint with
stout pectinate claw and 6 bristles (Fig. 5c).

Maxilla (Fig. Sf, g). Limb reduced. Dorsal margin of precoxale, coxale, and
proximal part of basale with fringe of hairs. Endite I with 3 ringed and 3 unringed
bristles; endite II with 2 ringed and 2 unringed bristles; endite III obscure, with
| proximal ringed bristle and 3 long, distal, ringed bristles. Coxale with | hirsute
dorsal bristle. Basale with | lateral ventral bristle, 1 medial bristle near dorsal
margin, and | distal medial bristle. Exopodite short, with 3 long bristles (Fig. 5g).
Ist endopodial joint with hairs in rows, | alpha-bristle and | beta-bristle; 2nd
endopodial joint with long hairs, 5 ringed bristles, and 2 bristles with rings only
distally (the latter 2 bristles stouter than others).

Fifth limb (Fig. 6a—d): Epipodial appendage with 48 bristles. Endite I with |

VOLUME 95, NUMBER 4 805

unringed and 2 ringed bristles; endite IJ with | unringed and 3 ringed bristles;
endite III with | short unringed bristle and 6 longer ringed bristles. Ist exopodial
joint with 3 ringed bristles and 3 unringed finger-like bristles (Fig. 6b). 2nd exo-
podial joint with 4 ringed bristles and | or 2 long, unringed, finger-like bristles
(Fig. 6c). 3rd exopodial joint with 3 slender bristles on inner lobe and 2 stout
hirsute bristles on outer lobe (FIg. 6d). 4th and Sth joints fused, hirsute, with
total of 5 bristles.

Sixth limb (Fig. 5h): Endite I and II each with 3 or 4 bristles (some bristles
broken on endite I of illustrated limb); endite III with 4 bristles; endite IV with
3 bristles. End joint with hairs on medial and lateral surface and 7 bristles (anterior
4 bristles with short marginal spines) following bristle with long hairs except near
spinous tip; posterior 2 bristles hirsute. Ventral margin of end joint linear but
bases of posterior 3 bristles on small pedestals. Two subequal bristles with long
marginal hairs in place of epipodial appendage.

Seventh limb (Fig. 51): 4 short widely spaced bristles in proximal group, 2 on
each side, each bristle with 24 bells, some bristles with marginal spines; 4 long
spinous bristles in terminal group, 2 on each side, each bristle with 3-5 bells.
Terminus consisting of comb with 9 alate teeth opposite 2 recurved spinous teeth.

Furca (Fig. 4h): Each lamella with 9 claws; claws 1, 2, 4, 5 stout primary claws;
claws 3, 6—9 small secondary claws; claws 1 and 2 with abundant slender teeth
along posterior margin (more numerous than on adult female or juvenile male);
claws 4 and 5 also with posterior teeth; anterior margin of lamella with long hairs
near base of claw | of right lamella; claw | or right lamella anterior to claw | of
left lamella. Teeth on main claws not shown on illustrated limb. Left lamella of
USNM 158489 aberrant in having claws 1, 2, 4, 5 and 6 main claws, and claws
3, 7-10 secondary claws.

Bellonci organ (Figs. 41, 6e): Elongate, with short broad segment near middle,
cylindrical distal part, and rounded tip.

Eyes: Medial eye bare, with black pigment (Figs. 41, 6e): Lateral eye well
developed, about same size as medial eye, with black pigment (Figs. 4f, i, 6e);
ommatidia large, but number difficult to determine because of pigment (14 visible,
but none shown in illustrations).

Upper lip (Figs. 41, 6e): Anterior process present but weaker and smaller than
on A-! male or adult female.

Copulatory organ: Consisting of short lobes, some with bristles.

Y-sclerite and posterior of body: Similar to that of adult female.

Comparisons.—The new species A. lizardensis differs from A. neali Kornicker
and Caraion, 1978:66 in that the carapace bears a distinct caudal process; also,
furcal claws 1, 2,3, 5 of A. neali are primary claws, whereas claw 3 is a secondary
claw on the furca of A. lizardensis. The furca of A. lizardensis does not have a
secondary claw between 2 sets of primary claws as on A. polychelata Kornicker,
1958:237.

Acknowledgments

My thanks to Peter N. Slattery, Moss Landing Marine Laboratory, Moss Land-
ing, California for collecting the specimens studied herein, and to Anne C. Cohen,
Smithsonian Institution, for commenting on the manuscript.

806 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Literature Cited

Kornicker, Louis S. 1958. Ecology and taxonomy of recent marine ostracodes in the Bimini area,
Great Bahama Bank.—Publications of the Institute of Marine Science (The University of Tex-
as) 5:194-300, 89 figs., | map.

, and Francisca Elena Caraion. 1978. West African myodocopid ostracoda (Sarsiellidae,
Rutidermatidae).—Smithsonian Contributions to Zoology, 250, 110 pp., 59 figs., 33 pls.
Poore, Gary C. B., and Sebastian Rainer. 1974. Distribution and abundance of soft-bottom molluscs
in Port Phillip Bay, Victoria, Australia.—Australian Journal of Marine and Freshwater Re-
search 25:371-411.

, Sebastian Rainer, R. B. Spies, and E. Ward. 1975. The zoobenthos program in Port Phillip
Bay, 1969-73.—Fisheries and Wildlife Paper, Victoria 7:78 pp.

Department of Invertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
95(4), 1982, pp. 807-825

DESCRIPTION OF MOOREONUPHIS JONESI, A NEW
SPECIES OF ONUPHID POLYCHAETE FROM SHALLOW
WATER IN BERMUDA, WITH COMMENTS ON
VARIABILITY AND POPULATION ECOLOGY

Kristian Fauchald

Abstract.—Mooreonuphis jonesi, new species (Polychaeta: Onuphidae) is de-
scribed from shallow water sandy bays on Bermuda. The variability of morpho-
logical features is discussed. Larvae and juveniles are described and selected
features of the population ecology of the species are discussed. The total length
and numbers of setigers of any specimen are shown to be correlated with mea-
surements obtainable on any incomplete specimen. This makes it possible to
calculate the total number of segments and the total length for all specimens of
a collection and thus, size-frequency distributions can be based on all specimens
sampled, rather than just on complete ones.

Shallow-water polychaetes from Bermuda have been reported by a series of
authors, including Webster (1884) and Verrill (1900). The fauna is reviewed in
- detail by Jones and Gardiner (in press). Early authors did not report any members
of the family Onuphidae from shallow water. In contrast, Hartman (1965), and
Hartman and Fauchald (1971) reported six species from slope and abyssal depths
off Bermuda; most of these species are limited to deep water areas.

The material studied by Jones and Gardiner (in press) is the same as that
reported upon here.

The generic definitions and terminology are as defined in Fauchald (1982). All
calculations follow the procedures outlined in Sokal and Rohlf (1969).

Most of the material is deposited in the collections of the National Museum of
Natural History, Smithsonian Institution (USNM), and paratypes have been de-
posited in the Allan Hancock Foundation, University of Southern California
(AHF), The Australian Museum, Sydney (AMS), British Museum (Natural His-
tory), London (BMNH), Zoologisches Museum und Staatsinstitut, Hamburg
(ZMH), and Zoological Institute, Academy of Sciences, Leningrad (ZIL).

Mooreonuphis jonesi, new species
Figures 1-8, Tables 1-4

Material examined.—Bailey’s Bay, inner bay near Fractious Street, inshore,
1.2 m depth, sand, 29 Aug 1981, coll. M. L. Jones (10 paratypes, USNM 72789);
Boaz Island, outer side of SW end of Grey’s Bridge, bare sand and sand overlain
with thick algal mat, 1.2 m depth, 6 Sept 1981, coll. M. L. Jones (13 paratypes,
USNM 72790); Boaz Island, outer side of SW end of Grey’s Bridge, Thalassia-
substrate, 1 m depth, 6 Sept 1981, coll. M. L. Jones (2 paratypes, USNM 72791);
Southeastern base of Causeway (Blue Hole side), sand, 14 Aug 1975, coll. M. L.
Jones (10 paratypes, USNM 72792; 2 paratypes, BMNH ZB 1982:43-44; 2 para-
types, AHF Poly 1373; 2 paratypes, AMS W 19107); Southeastern base of Cause-

808 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

=

=

J

Fig. 1. Mooreonuphis jonesi: a, Third parapodium, anterior view (paratype, USNM 72788); b,

Pectinate seta, median parapodium (holotype, USNM 72787); c, Anterior end, dorsal view; d, Anterior

end, lateral view; e, Aciculum, 14th parapodium; f—g, Pseudocompound hook, 4th parapodium; h-i,
Pseudocompound hook, Ist parapodium; j, Compound spiniger, 7th parapodium.

way (Blue Hole side), Thalassia, 14 Aug 1975, coll. M. L. Jones (6 paratypes,
USNM 72793); Coney Island, north side in fairly sheltered bay, clean white to
dark grey sand, | m depth, | June 1976, coll. M. L. Jones (6 paratypes, USNM
72794; 2 paratypes, ZMH P 17144; 2 paratypes, ZIL); Dennis’ Hideaway (restau-
rant on Smith’s Sound, Governor’s Island), clean sand, intertidal, 30 May 1976,
coll. M. L. Jones (2 paratypes, USNM 72795); Dennis’ Hideaway, clean sand,
50 cm depth, 30 May 1976, coll. M. L. Jones (45 paratypes, USNM 72796);
Dennis’ Hideaway, Thalassia, 50 cm depth, 30 May 1976, coll. M. L. Jones (7
paratypes, USNM 72797); Devonshire Bay, 150 m transect from 1.2 m depth to

VOLUME 95, NUMBER 4 809

ay OO

CCS Tes

Fig. 2. Mooreonuphis jonesi: a, Maxillae in dorsal view (paratype, USNM 72788); b, Maxillae of
13-setiger larva (paratype, USNM 72796).

shore, sand, 6 Sept 1981, coll. M. L. Jones (49 paratypes, USNM 72798); Ferry
Reach, adjacent to west side of dock at Bermuda Biological Station, Thalassia-
roots and substrate, | m depth, 2 Sept 1981 (2 paratypes, USNM 72799); Flatt’s
Inlet, northeast shore between Harrington Sound and Gibbet Island, west of
Flatt’s Bridge, sand, | m depth, 31 May 1976, coll. M. L. Jones (Holotype, USNM
72787; 17 paratypes, USNM 72788); Hungry Bay, sand, 16 Aug 1975, coll. M. L.
Jones (57, USNM 72801); Hungry Bay, clean sand, many rocks, |.2—1.5 m depth,
27 May 1976, coll. M. L. Jones (4, USNM 72802); Hungry Bay, outer half, mud-
sand substrate, 16 Nov 1979, coll. M. L. Jones (2, USNM 72803); Hungry Bay,
inner half, mud-sand substrate, 16 Nov 1979, coll. M. L. Jones (2, USNM 72804);
Hungry Bay, near beach, mud-sand substrate, 20 cm depth, 16 Nov 1979, coll.
M. L. Jones (174, USNM 72800); Hungry Bay, transect along the length of the
bay from 1.2 m depth to shore, outer 4, clean fine white sand, 5 Sept 1981, coll.
M. L. Jones (4, USNM 72805); Hungry Bay, transect along the length of the bay
from 1.2 m depth to the shore, middle 14, grey fine sand, 5 Sept 1981, coll. M.
L. Jones (2, USNM 72806); North Ireland Island, small bay on inner side, west
and outside the naval base breakwater, bare sand, | m depth, 6 Sept 1981, coll.
M. L. Jones (7, USNM 72807); North Ireland Island, small bay on inner side,
west and outside the naval base breakwater, mixed grass-bed, 6 Sept 1981. coll.

810 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Mooreoruphis jonesi: a, 5-setiger larva, lateral view (USNM 72813); b-c, Compound hooks,
first setiger of 13-setiger larva (paratype, USNM 72794); d, 8-setiger larva (paratype, USNM 72793);
e, Jaw-apparatus, ventral view, 8-setiger larva (paratype, USNM 72793); f-g, Compound hooks, Ist
setiger (USNM 72813); h-i, Compound hooks, Ist setiger, 13-setiger larva (USNM 72800).

M. L. Jones (26, USNM 72808); South Ireland Island, inner side of northeast end
of Grey’s Bridge, associated with rocks in subtidal, 31 Aug 1981, coll. M. L.
Jones (2, USNM 72809); South Ireland Island, inner side of northeast end of
Grey’s Bridge, bare sand, 31 Aug 1981 (14, USNM 72810); Spanish Point, small
boat harbor, near sunken barge breakwater, rocks and under rocks, 29 May 1976,
coll. M. L. Jones (17, USNM 72811); Spanish Point, small boat harbor, near
sunken barge breakwater, bare sand and rock, | m depth, 29 May 1976, coll. M.
L. Jones (6, USNM 72812); Spanish Point, small boat harbor, near sunken barge
breakwater, Thalassia, 29 May 1976, coll. M. L. Jones (6, USNM 72813); To-
bacco Bay, sand, 15 Aug 1975, coll. M. L. Jones (7, USNM 72814); Tobacco
Bay, Thalassia, 15 Aug 1975, coll. M. L. Jones (1, USNM 72815); Whalebone
Bay, sand, |! Aug 1975, coll. M. L. Jones (11, USNM 72816); Whalebone Bay,
sand, 14 Aug 1975, coll. M. L. Jones (21, USNM 72817).

Description of holotype.—The holotype is a complete specimen with I11 seti-
gers, 17.40 mm long and 0.64 mm wide at the widest. The anterior end (Fig. Ic,
d) including the first 15 setigers is cylindrical. The rest of the body is dorsally
flattened and convexly curved ventrally. The body is pale pink with a light orange
spot at the base of each ceratostyle. The anterior end of the body is only slightly

VOLUME 95, NUMBER 4 811

| Median occipital antennae
|

25

20 Start of branchiae
Digitiform postsetal lobes

15
tt Start of subacicular hooks
10

C-B HB DBS SWE) Wi 1975

Fig. 4. Range, mean and standard deviations for selected variable morphological features for
material collected in 1975. The samples include: C-B, Causeway-Blue Hole; HB, Hungry Bay; TB,
Tobacco Bay; WBI and WBII, two subsamples from Whalebone Bay. Ranges are indicated as single
Vertical lines, standard deviations as boxes, and means as horizontal lines.

narrower than the widest part of the body which is at setigers 15—20. The posterior
end tapers abruptly and the pygidium carries a pair of anal cirri which reach
setiger 15 from the posterior end when folded forward.

The prostomium (Fig. Ic, d) is a small rounded lobe. The triangular frontal
palps are about one-third as long as the prostomium. A pair of small, black eyes
are at the base of the inner lateral occipital antennae. The occipital ceratophores
are about as long as the length of the prostomium. Each ceratophore has 5 rings
of which the 4 basal ones together are less than half the length of the ceratophore.
The outer lateral occipital antennae reach setiger 2; the inner lateral antennae

812 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

10

Median occipital antennae

fp

2a Start of branchiae

15 an
Digitiform postsetal lobes
10

Start of subacicular hooks

CI DH FI HB Se 1976

Fig. 5. Range, mean and standard deviation for selected variable morphological features for ma-
terial collected in 1976. The samples include: CI, Coney Island, DH, Dennis’ Hideaway, FI, Flatt’s
Inlet, HB, Hungry Bay, SP, Spanish Point. Ranges are indicated as single vertical lines, standard
deviations as boxes, and means as horizontal lines.

reach setiger 9 and the median antenna reaches setiger 6. All ceratostyles are
tapered. The peristomial ring is about as long as the length of the prostomium
and the slender peristomial cirri reach the base of the inner lateral occipital an-
tennae.

VOLUME 95, NUMBER 4 813

10
Median

eccipital antennae

25

20 5 Start of branchiae

q
|

| o ¢ eee ee
tf

Start of

==
=]
ce
Es
=.

subacicular hooks

BB BI DB FR HB NII SII 198]

Fig. 6. Range, mean and standard deviation for selected variable morphological features for ma-
terial collected in 1981. The samples include: BB, Bailey’s Bay, BI, Boaz Island, DB, Devonshire
Bay, FR, Ferry Reach, HB, Hungry Bay, NII, North Ireland Island, SII, South Ireland Island. Ranges
are indicated as single vertical lines, standard deviations as boxes, and means as horizontal lines.

The first 3 pairs of parapodia (Fig. la) have projecting parapodial bases; each
has a distally rounded acicular lobe, a presetal lobe that follows the outline of
the acicular lobe, and a digitiform postsetal lobe. The first pair of parapodia are
no larger than the next following ones and do not project forward any further
than to the bases of the outer lateral occipital antennae. Digitiform postsetal lobes
are distinct in the first 14 setigers; ventral cirri are cirriform in the first 4 setigers
and are replaced by pad-shaped glands from setiger 5.

By setiger 5 the parapodial bases have been completely reduced and the parapo-
dia are sessile on the body-wall; the acicula project obliquely dorsally from the
body-wall near the ventral nerve chord.

Simple, strap-like branchiae are present from setiger 29 to setiger 85. Where
best developed, at setigers 40-50, each branchia is flattened and reaches beyond
the dorsal mid-line.

814

40

e0

20

]

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Total number of setigers Ro calelemane
ng

198]

1981

WO? N77
Pie WH
O75 WS

40

All specimens All specimens

30

20

10 41-50 HW-90 WeISO 16i)170 2ONZIO BAi-950 1-5 Me D5 AVeas 61-65

VOLUME 95, NUMBER 4 815

Setal array includes limbate and pectinate setae, pseudocompound and suba-
cicular hooks and larval-type hooks in addition to the acicula. All parapodia are
supported by a pair of acicula. Acicula in most setigers taper evenly to pointed
tips. Each aciculum in setigers 8-16 (Fig. le) is subdistally expanded into a large,
flattened pad; the tips remain tapered. Limbate setae are present in all setigers,
but are most common in the anterior third of the body. Pectinate setae (Fig. 1b)
are present from setiger 6; each is distally transverse or slightly oblique and has
11 teeth. Tridentate pseudocompound hooks (Figs. 1f—i) are present in the first 4
setigers. The lowermost tooth may be reduced to just a small knob on the side
of the hook and may be very difficult to see. Compound spinigers (Fig. 1j) are
present as an inferior fascicle in setigers 5—13. In setigers 4 and 5 a single, slightly
enlarged hook is present in the middle of each setal fascicle; these hooks do not
resemble the large hooks present in other onuphids, but closely resemble the
hooks present in larval. They are here considered remnant larval hooks. Bidentate
subacicular hooks are present from setiger 14.

The maxillary apparatus (seen in dissection in one paratype from Flatt’s Inlet,
USNM 72788, Fig. 2a) is poorly chitinized; the formula is 1+1, 9+10,6+0, 10+9
and 1+1.

The tubes are about twice as long as the contained specimens, narrow and
cylindrical in shape and have a tough inner lining covered externally by large
sand-grains and shell-fragments.

Mooreonuphis jonesi resembles M. veleronis (Fauchald, 1980:807—809, table 2)

‘in that these two are the only species known in the genus to have expanded
acicula in some anterior setigers; in all other species in the genus (reviewed by
Fauchald 1982:55—64) the acicula taper evenly to the tip in all setigers. Moor-
eonuphis jonesi differs from M. veleronis in that the expanded acicula are present
in setigers 8-15 in the former and in setigers 4—8 in the latter. Ventral cirri are
cirriform in four setigers in M. jonesi and in three in M. veleronis; digitiform
postsetal lobes are distinct in the first 14 setigers in M. jonesi and in the first five
setigers in M. veleronis. The occipital ceratophores are ringed with about five
rings in M. jonesi and are smooth in M. veleronis.

Mooreonuphis jonesi is known from several sandy beaches at Bermuda in shal-
low subtidal, sandy environments.

Etymology.—The species is named for Dr. Meredith L. Jones who collected
the material in Bermuda and who has followed my study of this extensive material
with more than casual interest.

Brood-chambers.—Larvae of M. jonesi are brooded in a chamber formed from
part of the tube. The brood-chamber is found some distance from the mouth of
the tube; it appears to be situated at the sediment-line where the adult tube
projects out from the sediment. The oval brood-chamber is usually visible from
the outside of the tube as a slight thickening of the wall of the tube. The chamber
projects into the tube and occupies about half the width of the tube. The larvae

—

Fig. 7. Size-frequency distribution diagrams. Left column, distribution of the total numbers of
setigers grouped in intervals of 10 setigers. Right column, distribution of the total length of the
specimens. Lower right diagram shows the frequency for the whole material combined.

816 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Total number of setigers Total length

Boaz Island

Devonshire Bay 198]

North Ireland Island

1981
| South Ireland Island 198]
Hungry Bay OA

Coney Island 1976
Flatts Inlet 1976

Causeway - Blue Hole NOTS

Hungry Bay 1975

Whalebone Bay

| | i
1-10 AES BIO VIO Wel170 20-210 24i-250 Pes eM EIST a lolls®:3

Fig. 8. Size-frequency distribution diagrams for selected individual stations. For further expla-
nation see legend for Figure 7.

VOLUME 95, NUMBER 4 817

Table 1.—Summary statistics for all specimens studied. Brooded specimens excluded.
Morphological feature Range Mean SD v/m n

Occipital antennae

Outer lateral reach [l= 1.09 97) .08 328

Inner lateral reach 4-14 Tals 1.58 “35 SIS)

Median reaches 2-10 5.63 1.41 5) 309

# of rings 0-6 S528) 81 IP 317
Branchiae

First present on setiger # 16—40 PANG 2.84 38 294

# of filaments | Invariant 294

Cirriform ventral cirri present

to setiger # 2-4 3.94 28 .02 328
Digitiform postsetal lobes

distinct to setiger # 4-19 13.58 1.88 .26 328
Pseudocompound hooks present

to setiger # 2-4 3.88 35 .03 328
Compound spinigers

First present on setiger # 4-5 4.90 25) .02 328

Last present on setiger # 9-17 15232 1.74 DB hie 328

Subacicular hooks first present
on setiger # 9-17 13.62 1.67 Pl ily

are oriented along the long axis of the chamber, usually about five to six deep
and in about 10 layers, so that about 50-60 larvae fill the chamber. The chamber
is subdivided internally by clear membranes so that each layer of five-six larvae
is isolated from the layers above and below it. The chamber is a distinct, ovate
structure, in contrast to the brood-chamber in Kinbergonuphis simoni which con-
sists of a half-cylinder stretching along about half the length of the tube in that
species (Fauchald, personal observation). The brood-chamber is a single large
unit, in contrast to the numerous small, lateral chambers found in Kinbergonuphis
notialis (Hartman 1967a, b).

Description of larvae.—TYhe brood-chambers contained developing early em-
bryos in addition to larvae in various stages of development. Eggs found in the
body-cavity of the adults measure about 175 micrometers on the average. The
eggs are somewhat flattened-ovate and the maximum long diameter is about 190
micrometers. Early embryos found in the brood-chambers measure about 200
micrometers in diameter. The massive amount of yolk-material present in each
egg had been too poorly preserved to allow a detailed description of the early
embryos. The larvae, however, are well preserved and are briefly described be-
low.

A five-setiger larva (Fig. 3a) is about 320 micrometers long and 175 micrometers
wide. It is essentially ovoid with a small, but distinct prostomium. All five oc-
cipital antennae are present as short, cylindrical, distally bluntly rounded rudi-
ments; the smooth ceratophores are distinct. The median occipital antenna is
usually best developed, but does not reach beyond the posterior edge of the
prostomium. The frontal palps are visible only as bulges on the anteroventral side
of the prostomium; they cannot be seen from the dorsal side. The peristomial

818 PROCEEDINGS OF THE BIOLOGICAESSOCIETY OF WASHINGTON

Table 2.—Correlations between paired morphological features from the different localities. The
features include: A, total number of setigers present; B, total _jength and E, relative size of each
segment, measured as the total length divided by the numbers of-segments, multiplied by 100.

Numbers of

complete
Locality specimens A-B A-E B-E
Bailey’s Bay 2 — = =
Boaz Island 4 80 .80 —
Causeway-Blue Hole S) 98 .93 —
Coney Island 5 96 91 —
Dennis’ Hideaway 8 — — =
Devonshire Bay 19 94 .80 —
Ferry Reach 3) 99 299 —
Flatt’s Inlet 6 92 78 —
Hungry Bay, 1975
adults 8 92 .85 —
juveniles 13 89 — WH
Hungry Bay, 1976 1 — — —
Hungry, Bay, 1979 20 82 — 88
Hungry Bay, 1981 2 — — —
North Ireland Island 11 94 — 86
South Ireland Island 8 94 .80 —
Spanish Point 5) — — —
Tobacco Bay — — — —
Whalebone Bay 8 .99 91 —

ring has not been differentiated from the remainder of the body and peristomial
cirri are absent. The first pair of parapodia has distinct ventral and dorsal cirri
as well as distinct, digitiform postsetal lobes; all other parapodia are blunt, short
paddles, decreasing in size and degree of differentiation posteriorly. A pair of
short, ovoid anal cirri are present. Setae include short, slender capillary setae
and compound, bidentate hooks (Figs. 3f—g) with both the distal end of the shaft
and the proximal end of the appendage finely dentate. A rudiment of the jaw-
apparatus is visible as a flattened indistinct area of sclerotinized material at the
junction of the prostomium to the rest of the body. The whole interior of the
larva is filled with a mass of yolk so the digestive tract is completely undiffer-
entiated. Muscles associated with the parapodia are well developed, and from
the position of the parapodia and setae it appears that the larvae at this stage are
able to crawl around in the brood-chamber.

An eight-setiger larva (Fig. 3d) is about 500 micrometers long and 135 micro-
meters wide. It is still oval with a large, bulging yolk-mass in the middle of the
body, but has elongated considerably since the five-setiger stage. It differs from
the five-setiger larva in that the median and inner lateral occipital antennae are
now distinctly digitiform; the blunt outer lateral antennae are still cylindrical. The
first parapodia are as in the five-setiger larvae, but dorsal cirri are now distinct
in the first 3 setigers. The anal cirri have elongated and are now about as long as
the inner lateral occipital antennae and the jaw-apparatus has differentiated. A
pair of rod-shaped, rather irregular mandibles (Fig. 3e) is supported by a pair of
separate mandibular carriers. The mandibular carriers are separate from each
other and from the mandibles. The 2 pairs of maxillary carriers are free from

VOLUME 95, NUMBER 4 819

Table 3.—Regression constants for each sample. A, total number of setigers; B, total length and
E, relative size of each segment.

Locality Regression a b
Boaz Island A-B —19.94 44
A-E —2.48 .26
Causeway-Blue Hole A-B —9.28 px)
A-E +2.10 ill
Coney Island A-B — 19.69 42
A-E —1.88 oD)
Devonshire Bay A-B = 2S LtS 58
A-E +1.74 26
Ferry Reach A-B =55). 10 69
A-E — 16.90 33
Flatt’s Inlet A-B —18.41 35
A-E (0; 1l7/ 22
Hungry Bay, 1975, adults A-B —23.24 37
A-E =§.39 21
juveniles A-B —0.19 10
B-E +4.32 a4
Hungry Bay, 1979 A-B = 119 03 45
B-E + 112.39) 46
. North Ireland Island A-B =).0z 38
B-E + 14.47 37)
South Ireland Island A-B —9.25 32
A-E = 7/8) 14
Whalebone Bay A-B —8.52 nes
A-E +3.78 11

———————_

each other. A single maxilla is present on the right side of the animal (seen on
the left in the drawing which shows the apparatus in ventral view). Two maxillae
are present on the left side. The first pair of maxillae, which are similar to each
other, consists of a series of separate, adjoining pieces, each of which carries a
tooth. The tooth of the distal piece is a large, curved hook. Maxillae I basally
are supported by a pair of carriers which are fused in the midline of the apparatus
and continuous on the dorsal side with the lightly sclerotinized lining of the buccal
cavity. Left maxilla II consists of two pieces. The basal piece is fused to its
carrier and is finely dentate marginally. The distal part of maxilla Hl consists of
a large, thickened hook, which has two or three teeth on the inner margin. The
carrier of maxilla II is free from the carriers of the first pair of maxillae. The left
maxilla I is distally covered by a folded, sclerotinized piece which is produced
distally into a pair of blunt, triangular teeth.

At about the ten-setiger stage each of setigers 6-8 develop a pair of large,
flattened, marginally ciliated branchiae. A 13-setiger larvae is about 675 micro-
meters long and 140 micrometers wide; its body is an elongated oval and all par-
apodia, except the last few, project laterally from the body. The yolk-mass is
nearly or completely obliterated and the gut-lining is distinct, at least through the

820 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 4.—Variability in size of adults from various localities. The columns are: 1, Locality; 2,
Numbers of complete specimens; 3, Total number of specimens; 4, Mean number of setigers; 5,
Standard deviation of mean number of setigers; 6, Variance to mean ratio of mean number of setigers;
7, Mean total length; 8, Standard deviation of mean total length; 9, Variance to mean ratio of mean
total length; 10, Mean length per setiger (x 100); 11, Standard deviation of mean length per setiger;
12, Variance to mean ratio for mean length per setiger; 13, Calculated values included: +, Calculated

values excluded: —. No complete specimens were available from Tobacco Bay.

1 re Vi 4 5 6 ai 8 9 10 li 12°) 7s
Bailey’s Bay De (OY 155) 1.41 .01 559.40 | 2.76 -.13 32.63. 4.98 S276" =
Boaz Island Ae 15 12220 16:04 2°29. 29743"~ 7.447 1-88 265830 4s 34 ee
Causeway-Blue Hole 9 20 118.15 36.64 11.36 17.83 8.36 3.92 15.17 3.99 1.05 +
Coney Island 5 10)... 100107" 37-60) 147 1226.99 9132-. 3322) 23712 eA Seen Oomme
Dennis’ Hideaway 8 (iv) 100-38 41291 17-33) 25,03. tee S05) 20F80N ese olae
Devonshire Bay 19. 40) \104:57 16.24 2.52 431.43) | 9993.05 295 4 83a Ome
Ferry Reach se ZO 1028 82 \7 32.90) >, Fn Se eS 20 e598, 3043 eee OM
Flatt’s Inlet 7 V4") 96.791 17.44 “3 .Se815.77._ Sv4OaeS9) Seo 93.64 eee Oonmne
Hungry Bay, 1975 21 35 82.97 36.43 16.00 11-59 8.485 -6:207 12 e469 loo

Hungry Bay, 1976 1 (4) 58 — — 6.72 — — 23.82 — —_—- -
Hungry Bay, 1979 20° 29 9122.93 16:85: 42.315 935.29) 1OM0N 2589" 29:03) 430 Ose
Hungry Bay, 1981 3° ©) 96:67 920.60! + 4.39120.99 8.48" 3.43927. 15 213537 eeo Some
North Ireland Island 11 33 126.00 47.04 17.56 32.51 15.61 7.50 26.49 5.90 1.31 +
South Ireland Island 8 16 103.44 28.97 8.11 23.98 9.58 3.83 22.45 4.65 96 +
Spanish Point > (20) 74.20" 35:56) 17204510760" 828590728) baa nS eieS
Whalebone Bay 8 31 123797 46.35 17-43 921052 “iss? S16) e359 ee

+

first 4 or 5 setigers. Dorsal cirri are distinct in the first 10 setigers and ventral
cirri are usually distinct in the first 2 setigers. Setae are of the same kind as in
the smaller larvae (Fig. 3c, h), except in the first setiger where the compound
hooks have been exchanged for a number of simple, distally falcate hooks (Figs.
3b, 1). In addition, in setigers 5-8, a single, distally tridentate hook is present;
this hook resembles the hooks found medially in the fascicles in setigers 4-5 in
the adults, but is considerably smaller. The first jaw-apparatus, described above
for the eight-setiger larva, has been shed and in several specimens was seen
lodged between the remnants of the yolk-cells in the incompletely differentiated
posterior end of the digestive tract. A new jaw-apparatus (Fig. 2b) has been
developed. The mandibles are fused medially and the mandibular carriers are
fused to the cutting edges. The maxillary apparatus resembles the one found in
the adults, except that the joint between the maxillary carriers and the first max-
illae is still incomplete. Maxillae I are distally falcate, left maxilla I has 4 basal
teeth, the right maxilla I has 7 basal teeth. Left maxilla II has 10 teeth, the right
maxilla II has 11. Maxillae II are distally curved upwards so that maxillae I are
held inside the curve of maxillae II when the jaws are fully withdrawn. Maxillae
III has 12 teeth and is closely appended to maxilla II. Left maxilla IV is short,
gently curved and has 6 teeth of which the proximal and distal ones are indistinct;
right maxilla IV is strongly curved and has 10 distinct teeth along the margin.
No later larvae were found in the brood-chambers. The smallest specimen
found free-living had 17 setigers; it is 648 micrometers long and 153 micrometers
wide with parapodia. The peristomium and peristomial cirri are fully formed and
the digestive tract appears fully formed but empty. The smallest specimen found

VOLUME 95, NUMBER 4 821

with gut contents had 25 setigers. The larval branchiae, present on setiger 6-8 in
the brooded juveniles, are absent in all free-living forms and juveniles up to about
the 30-setiger stage are abranchiate. The adult branchiae develop over the next
growth period, so that when the specimens first become sexually mature (at about
the 70-setiger stage), they have the adult complement of branchiae.

Variability of late juveniles and adults.—TYhe variability of morphological fea-
tures used to characterize the species is indicated in Table 1 and in Figures 4—6
(tables showing the variability of specimens from each sample are available upon
request). Most specimens examined are adults with at least 60 setigers (Fig. 7
bottom) and the range of variation of each morphological feature reflects this
limited size range of the specimens. Late juveniles are sparsely represented and
the ranges for most variable morphological features reflect this fact. However,
the distribution of digitiform postsetal lobes in the sample from Hungry Bay, 1975
(Fig. 4) indicates the presence of juveniles. More generally, the presence of ju-
veniles in the material from Flatt’s Inlet (Fig. 5) and from South Ireland Island
(Fig. 6) lead to the rather wider range for most morphological features in these
two samples than in the other samples.

While late juveniles can be identified by their branchial distribution, size, and
number of setigers present, the juveniles usually develop the adult complement
of most other features very rapidly. However, any one of a number of features
may lag behind the others in development. For example, a medium-sized speci-
_men from Flatt’s Inlet had digitiform postsetal lobes only in four setigers; all
other features were well within the normal range for specimens of this size and
from that locality. In other cases, the subacicular hooks are present at segments
earlier than would be expected.

The standard deviations are limited in most cases, indicating that most mor-
phological features are fairly closely controlled in this species. A study of the
variance to mean ratios in the table indicate that the variability of any feature is
largely independent of the position along the length of the body of any given
feature. The variability shown may reflect the mixed nature of the sampled pop-
ulations. It appears likely that at least two generations are present in most sam-
ples. If it had been possible to separate the different cohorts by independent
criteria, the variability shown by each cohort probably would have been consid-
erably less. The species broods its juveniles and these apparently settle imme-
diately upon release from the parent tube. Thus, each sample will mainly contain
siblings and perhaps other close relatives, reducing the expected variability within
each locality.

Over the seven-year period between the first and the last samples, all ranges,
means and standard deviations have remained reasonably constant, with no in-
dication of change over time in any of the characters investigated (Figs. 4-6). In
most instances the range is clearly larger than the standard devation, indicating
that a single unimodal distribution is present. For certain characters, and for
certain sampling localities, the standard deviations are larger than the range,
indicating that for the feature in question a bi- or multimodal underlying distri-
bution may be present. The start of the subacicular hooks especially shows this
pattern. A bimodal distribution might be related to differences between year-
classes or between sexes, but insufficient evidence is available to resolve this
problem.

822 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

The start of the subacicular hooks is related to the last occurrence of the
digitiform postsetal lobes. Means and standard deviations co-vary in nearly all
samples. Both features represent a transition between the anterior end and the
median part of the body. The digitiform postsetal lobes are characteristic of the
anterior parapodia in which the parapodial bases are supported by projecting
acicula; the subacicular hooks are characteristic of the remainder of the body in
which the parapodia appear more as low, rounded ridges on the convex ventral
side of the body than as lateral projections. The subacicular hooks themselves
represent a support feature for the parapodia in a part of the body where the
acicula point dorsally and laterally, the subacicular hooks point ventrally and
laterally thus strengthening the convex ventral side of the animal. The subacicular
hooks also probably function as points d’appui to the tube-wall, leaving the an-
terior end of the body free to move in and out of the tube.

The start of the branchiae is independent of the two features discussed above.
Most commonly the branchiae start between setigers 19 and 25; only in one
instance is the mean above 25 (Flatt’s Inlet, Fig. 5). In complete specimens the
branchiae usually end about 25 setigers from the posterior end, leaving the last
setigers abranchiate.

The length of the occipital antennae is independent of the features discussed
previously. Figs. 4-6 show only the distribution of the median antenna. The outer
lateral antennae are very nearly constant in length throughout the whole material,
and the length of the inner lateral antennae is very closely tied to the length of
the median antenna. In nearly all specimens the inner lateral antennae reach one
segment further than the median antenna, or are as long as the median antenna.
In only one or two cases are the inner lateral antennae shorter than the median
antenna among the 500+ specimens for which this feature could be scored.

Some of the localities sampled appear to have characteristic signatures in terms
of the variability among the adults. For example, in specimens from Tobacco
Bay the branchiae start late as do the subacicular hooks. Hungry Bay samples
have a characteristic signature in that the subacicular hooks start relatively early
and the antennae are short. These signatures make it possible to predict what
values to expect for these features in future collections from the various beaches
in Bermuda. The signatures indicated a certain isolation between the beaches,
possibly combined with characteristic differences in ecological conditions.

Size-frequency distribution of adults. —Most samples of polychaetes comprise
a few complete specimens and numerous anterior, median and posterior frag-
ments. Most ecological workers consider presence to be indicated by the number
of head-ends only and reported numbers of specimens usually indicate the num-
bers of head ends seen. For most purposes this procedure is adequate, but if
some measure of the life-diagram is desired, a means must be found for calculating
the size of all specimens sampled. The problem lies in finding one or more feature
that can be measured on all specimens, complete or not, and which is sufficiently
highly correlated with the total numbers of segments (or the total length of the
specimens) so that it can be used as an estimate of the size of all specimens
sampled.

Usually the number of complete specimens is too small to allow the calculation
of correlation coefficients and regressions, so a study of the size-frequency dis-
tribution is impossible. The current material was very carefully collected and as

VOLUME 95, NUMBER 4 823

a result, in most samples an adequate number of complete specimens are present.
Generally, the total number of segments is highly correlated with the total length
of the specimens. Furthermore, the average length of each individual segment,
measured as the total length divided by the total number of segments (100), is
highly correlated with either of the two parent measurements. The average length
of each segment can be calculated for fragments as well as for complete speci-
mens, and, thus, one has a way of calculating the size of all incomplete specimens.
These calculations were done for the head-ends only; the median and posterior
fragments present must belong to one or another of the head-ends present; no
attempts were made at associating these bits with each other or with any of the
head-ends.

Table 2 reviews the correlations calculated and the correlation coefficients for
the regressions used are indicated. The regression equations are given in Table
3. Arbitrarily, no regressions were calculated unless the correlation coefficients
were higher than 0.75; in most instances the correlation coefficients for the regres-
sions used where higher than 0.80. For small samples, the calculation of both
coefficients and regressions are subject to huge errors, and the interpretation of
the results are here done in the form of an indication of trends, rather than of a
detailed numerical analysis.

Correlation coefficients and regression constants vary among samples (Tables
2-3), indicating that the growth patterns for this species vary from one bay to the
_ next on Bermuda. This variation is not surprising; it may be caused by slightly
differing conditions in terms of wave-action and food-supply in each cove and
these factors would be reflected in the growth pattern of the fauna of the coves.
As indicated above, the variation may also reflect slight genetic differences among
the populations and thus indicate a degree of isolation in the coves.

The size-frequency distribution for all specimens indicates that a specimen
grows to a normal upper limit of about 130 segments and that this number is fairly
precisely controlled. A linear plot of the length versus total numbers of segments
shows a distinct break in the curve at about 75 setigers indicating that the growth
pattern changes from a rapid increase in length to a more rapid increase in num-
bers of setigers. Inadequate numbers of specimens are available at any locality
to test this change in growth pattern and a summary plot for all specimens lacks
this distinct break in the curve, possibly because the change in growth patterns
happens at slightly differing sizes in the different bays. Interestingly, the smallest
sexually mature female found had 75 setigers.

The implications of these findings are that it appears that the measurement of
size in polychaetes is not as subject to variation as previously assumed; and that
at least some species with apparently large and variable numbers of segments, in
fact control the total numbers of segments produced fairly precisely. There are
no indications that anterior regeneration influences these calculations apprecia-
bly. Combining measurements of several different specimens show that a limited
number of anterior setigers, probably about 10, may be regenerated, and that the
necessary additional segments, if more are missing, are added posteriorly to get
the specimens back up to the usual adult complement of segments.

Table 4 reviews the mean measurements in size for the different localities. It
is worth noting that the standard deviations are narrower by an order of magnitude
for the measurements of the average length per segment. The size-frequency

824 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

distribution for specimens collected in the different years of sampling (Fig. 7)
shows clearly that each year has a different pattern. For example, 1975 was.
dominated by small specimens and 1979 by large specimens, whereas 1981 most
closely mimics the total size-frequency distribution of any of the years sampled.
It is worth noting that while the numbers of setigers show a clearly bell-shaped
pattern with a strongly peaked center, the distribution of the total length is skewed
strongly towards the smaller sizes, again indicating the different processes in-
volved with increase in size in these polychaetes. Individual samples taken the
same year also show considerable differences (Fig. 8). In 1975, which as a whole
was dominated by small specimens, most of these small individuals came in a
single sample from Hungry Bay; note also the difference between Hungry Bay
and Whalebone Bay in terms of the total numbers of setigers present. In Dev-
onshire Bay it is apparent that a single population was sampled; both length and
numbers of segments show clear bell-shaped distributions; in the same year, the
sample from North Ireland Island indicates the presence of more than one size
group of individuals.

Life-diagram pattern.—Samples were taken in November, May, June, August
and September. Sexually mature females were present in all months and brooded
larvae were present at all times, indicating that the species breeds the year round
at Bermuda. Reproductive activity appears evenly distributed in all months sam-
pled, so there is no evidence for a limited, or focused breeding season. The size-
frequency distribution of the adults indicate a single annual cycle with the animals
maturing after about a year of growth. Neither the total number of segments nor
the total length distribution show any trace of a bimodal distribution in the pop-
ulation. However, if one examines the size-distribution of the sexually mature
females a slightly different pattern emerges. The numbers are very small, only
two or three specimens in each category, so no statistical demonstration 1s pos-
sible, and the hypothesis may be entirely based on sampling error. Nevertheless,
the mature females are separated into four groups, one of small animals, at about
75 segments, one group at about 100 segments, the largest group, five specimens,
at about 125 segments and one single super female (the largest specimen collected)
has about 160 segments. This pattern might emerge if the species was living for
about five years, with only the females surviving for more than about three years.
It is difficult without sectioning to separate males from immature females, but
the few observations available show that no male larger than about 100 segments
has been found. The males are generally considerably narrower than the females
and some of the variability in size-measurements appears to be due to a slight
sexual dimorphism, but, as indicated above, the difference between immature
females and males is difficult to detect.

It is at this point not possible to demonstrate very accurately how long the
larvae are brooded in the tubes, nor the longevity of the species. As a working
hypothesis it appears likely that at least some specimens of the species live up
to five years, that all specimens become sexually mature at the end of the first
year and that the females reproduce at least once a year thereafter. Considering
the relatively low numbers of mature females present every season, it appears
likely that each female reproduces only once a year. A tenuous piece of sup-
porting evidence might be mentioned: three females with large brooded larvae in
their tubes also had nearly full-sized eggs in their body cavities, indicating that

VOLUME 95, NUMBER 4 825

perhaps the brooding period is fairly long, and certainly, that each female repro-
duces more than once.

Acknowledgments

I would like to thank Dr. Meredith L. Jones for permission to study the very
large material he collected in Bermuda and for his interest in and support of my
study.

Literature Cited

Fauchald, K. 1980. Onuphidae (Polychaeta) from Belize, Central America, with notes on related
taxa.—Proceedings of the Biological Society of Washington 93(3):797-829, 7 figures, 6 tables.

—. 1982. Revision of Onuphis, Nothria, and Paradiopatra (Polychaeta: Onuphidae) based upon
type-material.—Smithsonian Contributions to Zoology 356:1—109, 28 figures, 34 tables.

—. In press. Life-diagram patterns in benthic polychaetes.—Proceedings of the Biological
Society of Washington.

Hartman, O. 1965. Deep-water benthic polychaetous annelids off New England to Bermuda and
other North Atlantic areas.—Allan Hancock Foundation Occasional Papers 28: 1-378, 52 plates.

—. 1967a. Larval development of benthic invertebrates in Antarctic seas: early development
of Nothria notialis (Monro) and Paronuphis antarctica (Monro) in Bransfield Strait, Antarctic
Peninsula.—Proceedings of the Symposium on Pacific-Antarctic Sciences, JARE Scientific
Reports Special Issue 1:205—208.

—. 1967b. Polychaetous annelids collected by the USNS Eltanin and Staten Island cruises,

chiefly from Antarctic Seas.—Allan Hancock Monographs in Marine Biology 2:1—387, 51 plates.

, and K. Fauchald. 1971. Deep-water benthic polychaetous annelids off New England to

Bermuda and other North Atlantic areas. Part 2.—Allan Hancock Monographs in Marine

Biology 6:1—327, 34 plates.

Jones, M. L., and S. L. Gardiner. In press. Polychaeta.—Jn: Sterrer, W. (ed.): Marine fauna and
flora of Bermuda.

Sokal, R. R., and F. J. Rohlf. 1969. Biometry. The principles and practice of statistics in biological
research.—W. H. Freeman and Company, San Francisco, 776 pp.

Verrill, A. E. 1900. Additions to the Turbellaria, Nemertina and Annelida of the Bermudas, with
revisions of some New England genera and species.—Transactions of the Connecticut Acad-
emy of Arts and Sciences 10: 595-671, | plate.

Webster, H. E. 1884. Annelida from Bermuda.—Bulletin of the United States National Museum
25:305-327, plates 7-12.

Department of Invertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560.

826 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

BIOLOGICAL SOCIETY OF WASHINGTON
PROCEEDINGS

1060 Meeting—14 April 1982

The one hundred and third annual meeting was called to order by the President,
Raymond Manning, with twenty-five members present. The minutes of the
previous annual meeting were approved.

The reports of the various officers were presented and accepted. The Treasurer,
Leslie Knapp, stated that $38,215 is available in endowment funds at this time.
The applicable interest rate is 13.8%. The Society had $63,679 in 1980. The de-
crease in available funds is a result of the $3,000 expenditure for the Centennial
activities, and for the support of free pages. Actions have been taken recently to
increase the income. Rates were raised in 1981 to $18 for U.S. and $23 for foreign
subscribers. Page charges were increased to $40 per page. In addition, it has been
recommended that not as many free pages be provided during the coming year.
It is expected that prices will increase next year; Allen Press bills are expected
to increase as are the cost of paper and postal rates. The substitute editor, Stephen
Cairns, reported that the first issue of volume 95 has just been submitted to the
printer; it is the first to be printed in the broader format. Number 2 will be
submitted in July. Number 3 is two-thirds full and will be submitted in September.
Ninety-nine papers were submitted in 1981; to date, forty-five papers have been
submitted in 1982. Appreciation was expressed to Steve for the excellent work
he has performed in Editor Kensley’s absence.

The activities of the council in the past year were presented and discussed.
These include the following points. The Society is considering meeting with the
American Society of Zoologists in Philadelphia in December 1983. It has been
suggested that the terms of elected members be lengthened; this possibility is
under investigation. There is a proposal that advertisements of the Society be
prepared and used in order to assist in obtaining new subscriptions. It was rec-
ommended that the Biological Society of Washington publication be referenced
in more of the existing abstracts.

The officers were thanked for their efforts and the newly elected officers were
announced:

President, Paul J. Spangler

Vice President, David L. Pawson
Secretary, Catherine J. Kerby
Treasurer, Leslie W. Knapp

Councilors: Janet C. Gomon
Gordon L. Hendler
Richard S. Houbrick
Roy W. McDiarmid
James C. Tyler

Catherine J. Kerby
Secretary

INFORMATION FOR CONTRIBUTORS

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CONTENTS

Pterynotus xenos, a new species of muricid from off northern Jamaica (Mollusca: Gastropoda)
M. G. Harasewych

Istigobius hoesi, a new gobiid fish from Australia (Perciformes: Gobiidae)
Edward O. Murdy and John D. McEachran
Subspecies and geographic distribution of Black-mantle Tamarins Saguinus nigricollis Spix

(Primates: Callitrichidae) Philip Hershkovitz
Review of the genus Micrognathus Duncker (Pisces: Syngnathidae), with description of
M. natans, n. sp. C. E. Dawson

Taxonomy of Bufo venustus ee 1899 (Anura: Leptodactylidae) from central Chile
J. R. Formas and Alberto Veloso
Gymnodorvillea floridana, a new genus and species of Dorvilleidae (Polychaeta) from south-

eastern Florida Sam C. Wainright and Thomas H. Perkins
Description of Dactylokepon sulcipes n. sp. (Crustacea: Isopoda: Bopyridae) and notes on
D. caribbaeus | Daniel L. Adkison
Notalpheus imarpe: a new genus and species of snapping shrimp from western South
America (Decapoda: Alpheidae) Matilde Méndez G. and Mary K. Wicksten
South American freshwater needlefishes of the genus Potamorrhaphis (Beloniformes:
Belonidae) Bruce B. Collette
A revision of the genus Siphonosoma (Sipuncula) Edward B. Cutler and Norma J. Cutler

A new species of Euglandina from Peru (Gastropoda: Pulmonata: Spiraxidae)
Fred G. ‘Tienes
Description of a new cypridopsine genus (Crustacea: Ostracoda) from Campbell Island, with a

key to the Cypridopsinae K. G. McKenzie
Evoplosoma virgo, a new goniasterid starfish (Echinodermata: Asteroidea) from the Gulf of
Mexico Maureen E. Downey
Paraprovincialism: remnants of paleoprovincial boundaries in Recent marine molluscan
provinces etl Edward J. Petuch
A eunicid polychaete from a white smoker Kristian Fauchald

Curimatopsis myersi, a new curimatid characiform fish (Pisces: Characiformes) from Paraguay

Richard P. Vani

Alternochelata lizardensis, a new species of myodocopine ostracode from the Great Barrier

Reef of Australia (Rutidermatidae) Louis S. Kornicker

Description of Mooreonuphis jonesi, a new species of onuphid polychaete from shallow water
in Bermuda, with comments on variability and population ecology

Kristian Fauchald

639

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