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S14 OGT2

PROCEEDINGS

of the

Biological Society of
Washington

VOLUME 94

1981
Vol. 94(1) published 19 May 1981 Vol. 94(2) published 20 July 1981
Vol. 94(3) published 16 October 1981 Vol. 94(4) published 31 December 1981
WASHINGTON

PRINTED FOR THE SOCIETY

EDITOR

BRIAN KENSLEY

ASSOCIATE EDITORS

Classical Languages Invertebrates

GEORGE C. STEYSKAL THOMAS E. BOWMAN

Plants Vertebrates

Davip 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 1980-1981

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 94

Aldrich, John W. Geographic variation in white-winged doves with reference
to possible source of new Florida population ____________-__________________________-
Alvarino, Angeles. Spadella legazpichessi, a new benthic chaetognath from
Enewetak: Marshalllssland's: (. 22 25 Nice oe ese ee ee oe
Baker, H. R. A redescription of Tubificoides heterochaetus (Michaelsen) (Oli-
gochaeta:sLubitictdae):| 20a. area Tae ee nee rae ee a ese eg ce
Barnard, J. Laurens. Redescription of Iphiplateia whiteleggei, a New Guinea
marine Amphipod (Crustacea) 22-22 eee
Barnard, J. Laurens, and Charline M. Barnard. The amphipod genera Eobrolgus
and Eyakia (Crustacea: Phoxocephalidae) in the Pacific Ocean ____________________
Bayer, Frederick M. Pseudoplumarella echidna, a new species of Primnoid
octocoral from Queensland (Coelenterata: Octocorallia) ____._______________________
Bayer, Frederick M. On some genera of stoloniferous octocorals (Coelenterata:
Anthozoa), with descriptions of new taxa ___________________________-____--__-_--_-
Bayer, Frederick M. Key to the genera of Octocorallia exclusive of Pennatulacea
(Coelenterata: Anthozoa), with diagnoses of new taxa _____________________________-
Blake, James A. The Scalibregmatidae (Annelida: Polychaeta) from South
America and Antarctica collected chiefly during the cruises of the R/V Anton
Bruun, ORIN Hero, andvU SNS Elianin, 2.23 ee
Blake, James A., and Keith H. Woodwick. The morphology of Tripolydora
spinosa Woodwick (Polychaeta: Spionidae): an application of the scanning
electron microscope! toy Roly chaete SyStemmatics mae useeusees ass en en uyiale enue
Bowman, Thomas E. First description of the male opossum shrimp, Heteromysis
bermudensis bermudensis (Crustacea: Mysidacea) _________-----_-__--___-__-__-____-
Bowman, Thomas E. Calasellus longus, a new genus and species of troglobitic
Asellid from Shaver Lake, California (Crustacea: Isopoda: Asellidae) ____________
Brinkhurst, Ralph O. A contribution to the taxonomy of the Tubificinae (Oligo-
chaeta:. ‘Mubificidae) \icc20 ts aoe 8 ae a 2 Pie ee
Burreson, Eugene M. A new deep-sea leech, Bathybdella sawyeri, n. gen., n. sp.,
from thermal vent areas on the Galapagos Rift ______________________________________
Chapman, Peter M. A new species of Homochaeta (Oligochaeta: Naididae)
fromthe west coastofl- Canada; oc. SA ia eae ee
Chernoff, Barry, and Robert Rush Miller. Systematics and variation of the
Aztec Shiner, Notropis sallei, a cyprinid fish from Central Mexico ______________
Cohen, Daniel M. Saccogaster melanomycter (Ophidiiformes: Bythitidae), a new
fishespecies tromuthe: Cantbbean 2. 2 25 oe
Cohen, Daniel M. New and rare ophidiiform fishes from the eastern Atlantic:
Cananyslslandsito: the: Cape ols Goodshlope 2a sae esse
Cooke, William J., and Craig D. MacDonald. The puerulus and post-puerulus
of the Hawaiian spiny lobster Panulirus marginatus ___-----------------------------
Cressey, Hillary Boyle. Ceratocolax mykternastes, new species (Copepoda:
Bomolochidae) parasitic in the nasal sinus of Haemulon sciurus (Pisces:
Pomadasyidae) from) Belize oi 5 2 ike poe a eas ae tO) eae ee eee eee
Cutler, Edward B. A new species of Aspidosiphon (Sipuncula) from the western
Atlantic OCe an xi cert BOS RN arly tia re OL ia ci | etd ca OE
Dawson, C. E. Notes on west African pipefishes (Syngnathidae) with descrip-
Honvol-EnneacanipuUs, IW. PMs) 255. ce oe 8 ke es oe ae ee oe ne ae eee

641-651

107-121

564-568

1211-1218

295-313

622-630

878-901

902-947 |

1131-1162

352-362

458-461

866-872

1048-1067

483-49]

455—457

18-36

374-377

1085-1103

1226-1232

514-524

445-449

464-478

DeWitt, Hugh H., Paul A. Grecay, John S. Hacunda, B. Paul Lindsay, Richard F.
Shaw, and David W. Townsend. An addition to the fish fauna of the Gulf
olaViaineawithine cords oferane species ees ee ee eee ee

Downey, Maureen E. A new Goniasterid seastar, Evoplosoma scorpio (Echino-
dermata: Asteroidea), from the northeastern Atlantic ____-_.----------

Erséus, Christer, and Michael S. Loden. Phallodrilinae (Oligochaeta: Tubificidae)
from the east coast of Florida, with description of a new species of Adelo-
ONT IDIS oat 9s OO hp 8 SS eit oP ERE eS MSS A ACRE RLED ne eed Sd pe USL CT

Ewing, R. Michael, and Daniel M. Dauer. A new species of Amastigos (Poly-
chaeta: Capitellidae) from the Chesapeake Bay and Atlantic coast of the United
States with notes on the Capitellidae of the Chesapeake Bay ______________________

Farfante, Isabel Pérez. Solenocera alfonso, a new species of shrimp (Penaeoidea:
Solenocenidae)stromb these hilip pine Syn ee Ei ee ea esas)

Ferrari, Frank D. Oithona wellershausi, new species, and O. spinulosa Lindberg,
1950 (Copepoda: Cyclopoida; Oithonidae) from the mouth of the Pearl River,

Fritzsche, Ronald A. A new species of pipefish (Pisces: Syngnathidae: Micro-
SLATES ELO Hes A NUTR SCRE, NTT UY te DEE OR Re Ohta es ee ie
Froeschner, Richard C., and Libby Halpin. Heteroptera recently collected in the
RaveMountarmseinvAlaskitestent tesa IP ieee eT ASS I AU 8 UN
Gaston, Gary R., and David A. Benner. On Dorvilleidae and Iphitimidae (An-
nelida: Polychaeta) with a redescription of Eteonopsis geryonicola and a new
OS GMC CONG IRE eS tines at nn eRe iae) . UE aS TED. CC Ea sea, Wat
Gore, Robert H. Three new shrimps, and some interesting new records of deca-
pod Crustacea from a deep-water coral reef in the Florida Keys __________________
Grabe, Stephen A. Occurrence of Mysidopsis almyra Bowman, 1964 (Mysidacea)
in the Patapsco River Estuary (Upper Chesapeake Bay), Maryland, U.S.A. ______
Grogan, William L., Jr., and Willis W. Wirth. Anew American genus of predaceous
midges related to Palpomyia and Bezzia (Diptera: Ceratopogonidae) ___________-
Grygier, Mark J. Gorgoniscus incisodactylus, a new isopod of the epicaridean
superfamily Cryptoniscoidea, parasitic on an ascothoracican cirriped from
|S ENGI <a eS Ser as Ae «SUE A UANE AS Derailed Bes STE TERRE ON ceo aPR ETS py Ae!
Grygier, Mark J. Scalpelloniscus, a new genus for two species of cryptoniscid
isopods (Epicaridea) parasitic on abyssal stalked barnacles ________________________
Haman, D., K. L. Finger, and R. W. Huddleston. Bicornifera Lindenberg,
1965 emend. Keij, 1969 (Microproblematica) from the type Byram Marl (Oligo-
Cene) ke MiISSISSip pit. 20ee" anche RIS oe his te eR eal
Haman, Drew, and Richard W. Huddleston. Falsipatellina, a new name for
Pseudopatellina Kenawy and Nyiro, 1967 non Haque, 1960 (Foraminifer-
FUGUE) EAC) Meg PME AA SI ce 0 Ek ot pe =r ge vy Oe
Hastings, Philip A., and Stephen A. Bortone. Chriolepis vespa, a new species
oGobidifish trom the northeastern! Gulfof Mexico 222 a
Ho, Ju-shey, and Laurie E. McKinney. A new species of Eudactylina (Copepoda:
Eudactylinidae) parasitic on black shark from Chile .2.------- 8
Holt, Perry C. A resumé of the members of the genus Cambarincola (Annelida:
Branchiobdellida) from the Pacific drainage of the United States ________._________
Holt, Perry C. New species of Sathodrilus Holt, 1968, (Clitellata: Branchiob-
dellida) from the Pacific drainage of the United States, with the synonymy of
SOMOS VIRGINA VN OMS TT ees es es le Sen te i ed
Holthuis, L. B. Description of three new species of shrimps (Crustacea:
Decapoda: Canidea) ainom baci Cis lam Site eee saa tee ee eee ee

669-674

561-563

819-825

163-168

631-639

1244-1257

TN-173

423-426

76-87

135-162

863-865

1279-1305

122-134

1258-1270

696-698

378-379

427-436

745-752

675-695

848-862

787-800

Houbrick, Richard S. Anatomy of Diastoma melanioides (Reeve, 1849) with
remarks on the systematic position of the family Diastomatidae (Prosobranchia:
Gastropoda) io {os ae es = eg ea oe palais. Sete ee ee

Houbrick, Richard S. Systematic position of the genus Glyptozaria Iredale
(Prosobranchia: (Gastropoda) 2225. 2 hs he ete al

Huddleston, Richard W. Bernardichthys zorraquinosi, a new genus and species
of salmoniform fish from the late Cretaceous of Oregon ____________________________

Huddleston, Richard W. Comments on the taxonomic status of Tommotia
Missarzhevsky 1970 (Tommotiidae: Microproblematica) ____________________________

Huddleston, Richard W., and Drew Haman. Pflugella, new name for Tricellaria
Pflug, 1965 (Microproblematica), non Fleming, 1828 (Bryozoa) ______-__-_-_____-

Huddleston, Richard W., and Prabha Kalia. Shastrina, a new genus of hetero-
helicidiForaminifera trom! the Eocene of India sa aeeeu uns eees oe

Humes, Arthur G. A new species of Tegastes (Copepoda: Harpacticoida)
associated with a scleractinian coral at Enewetak Atoll ___._______________--_-______

Jara, Carlos G., and Maria T. Lopez. A new species of freshwater crab (Crus-
tacea: Anomura: Aeglidae) from insular south Chile ________________________________

Karnella, Susan J., and Ernest A. Lachner. Three new species of the Eviota
epiphanes group having vertical trunk bars (Pisces: Gobiidae) ____-______-________

Kimsey, Lynn Siri. The Cleptinae of the Western Hemisphere (Chrysididae:
Hymenoptera) iter k set Poo oT lee ah eee kh ee

Kornicker, Louis S. Range extension and supplementary description of Bathy-
conchoecia deeveyae (Ostracoda: Halocyprididae) ________-____-___________________-

Kornicker, Louis S., and Anne C. Cohen. The relative position of the left and
right lamellae of the furca in the order Myodocopida (Crustacea: Ostra-
COMA) - 2 cee eh end ROS: teens pyc AN ae ue od ht pati eR. er ery utara tl ke,

Lewis, Julian J. Caecidotea salemensis and C. fustis, new subterranean Asellids
from the Salem Plateau (Crustacea: Isopoda: Asellidae) ____________________________

Maciolek, Nancy J. A new genus and species of Spionidae (Annelida: Polychaeta)
fromthe montheand ;SouthyA thea tic pe ses esse re ag a

Maciolek, Nancy J. Spionidae (Annelida: Polychaeta) from the Galapagos Rift
PSCOMMEKIMALGVEMUS ss. ae 2 ede a Le

Manning, Raymond B., and David K. Camp. A review of the Platysquilla
complex (Crustacea: Stomatopoda: Nannosquillidae), with the designation of
{WO NeW EMEA eu: see ee SEN il 8 mena ied 2 ean

Manning, Raymond B., and C. W. Hart, Jr. Gonodactylus lightbourni, a new
Stomatopodicrustacean tiomi Bermuda 22 eee

Manning, Raymond B., and Marjorie L. Reaka. Gonodactylus siamensis, a new
Stomatopod crustacean from Mhailand) =) = eee

Marelli, Dan C. New records for Caprellidae in California, and notes on a
morphological variant of Caprella verrucosa Boeck, 1871 ___-----------------------

Mayes, Monte A., Daniel R. Brooks, and Thomas B. Thorson. Potamotry-
gonocotyle tsalickisi, new genus and species (Monogenea: Monocotylidae) and
Paraheteronchocotyle amazonensis, new genus and species (Monogenea:
Hexabothriidae) from Potamotrygon circularis Garman (Chondrichthyes: Pota-
Mothyeonidae) insnonthwestern! Brazil ee eee ne eee re eke ener

Messing, Charles Garrett. Reclassification and redescription of the comatulid
Comatonia cristata (Hartlaub) (Echinodermata: Crinoidea) _______________________-

Mikkelsen, Paula M. Studies on euphausiacean crustaceans from the Indian
River region of Florida. I. Systematics of the Stylocheiron longicorne species-
group, with emphasis on reproductive morphology __________-_---------_--_------__-

Vi

598-621

838-847

37-42

462-463

421-422

652-653

254-263

88-93

264-275

801-818 ©

1237-1243

739-744

579-590

228-239

826-837

591-597

708-712

479-482

654-662

1205-1210

240-253

1174-1204

Miller, Walter B. A new genus and a new species of Helminthoglyptid land
Snailssiromithe Mojave Desertot: California = sees as ones sande see a eee
Miller, Walter B. A new Greggelix (Mollusca: Pulmonata: Helminthoglyptidae)
{rom Bajar CalifornmiatS ur, Mexico ese tion tae. Tos eens ee ee ps et eve
Murdy, Edward O., Carl J. Ferraris, Jr., Douglass F. Hoese, and Roger C.
Steene. Preliminary list of fishes from Sombrero Island, Philippines, with
HINTS CMEC WAKE CONC Sy eee seen ree Ue Te See Oo ae eae I SS AE
Olson, Storrs L. A revision of the northern forms of Euphonia xanthogaster
CAvics ei lhraupid ac) ieee ta Wh. Me ONE SSE Et SS Pay ead et ane Rae ieee
Olson, Storrs L. A revision of the subspecies of Sporophila (‘‘Oryzoborus’’)
aneolensisa(AVeswe mbehizinac)\ ste See a ne oie a
Olson, Storrs L. Systematic notes on certain oscines from Panama and adjacent
ANCASH CAVE Stu ba SSEnILORMIES) asa s As ein Ei gs nS ere ee ee
Olson, Storrs L. The nature of the variability in the variable seedeater in Panama
(porophilaramericana ae Mbenizinae) meas lectin eee ey! ae
Pawson, David L., and Cynthia A. Gust. Holothuria (Platyperona) rowei, a
new sea cucumber from Florida (Echinodermata: Holothuroidea) ________________
Pawson, David L., and John E. Miller. Western Atlantic sea cucumbers of the
genus Thyone, with description of two new species (Echinodermata: Holo-
(i UU CONVO (SN 3 fe 5 WR Se eet lage Gare Re hg Eee yee gn Se oe eR
Pawson, David L., and John F. Valentine. Psolidium prostratum, new species,
from off the east coast of the U.S.A. (Echinodermata: Holothuroidea) _________-
Petuch, Edward J. A volutid species radiation from northern Honduras, with
notes on the Honduran Caloosahatchian secondary relict pocket __________________
Poore, Gary C. B., and Brian Kensley. Coralanthura and Sauranthura, two new
genera of Anthurideans from northeastern Australia (Crustacea: Isopoda: An-
CEENUITE Cl Ve) eee et sn PS Oe SA reir a ea ead Bs ny ek Si eae Rees yo RINE
Pyburn, William F. A new poison-dart frog (Anura: Dendrobatidae) from the
ORESE. OE SOUT NSS ISIin (OO) CoynMoyey ee eee ae
Pyburn, William F., and John D. Lynch. Two little-known species of Eleuthero-
dactylus (Amphibia: Leptodactylidae) from the Sierra de la Macarena, Colom-
Diep ee ot he ht Sere val Mur Pes me yeahs open te! BPA stots Rate ee aeareie fuer
Ramirez-Pulido, José, and Marie Claire Britton. An historical synthesis of Mexi-
CANnenammalianitaxOnOimy UA Ae week hh bee a0) Bahk nace ian 2 eee
Remsen, J. V., Jr. A new subspecies of Schizoeaca harterti with notes on
taxonomy and natural history of Schizoeaca (Aves: Furnartidae) _____-____-_____
Rosewater, Joseph, and Dietrich Kadolsky. Rectifications in the nomenclature
olesome Indo-Raciczittonmdac—ll See ee ee ee ee
Rozbaczylo, N., and J. C. Castilla. Australonuphis violacea, a new polychaete
(@nuphidae)mnomithe, southeast Racitic Ocean) 2 eee
Rutzler, Klaus, and Richard G. Bromley. Cliona rhodensis, new species (Porif-
era-wadromenida)) tromithe Mediterranean: y.2.) 2 a menn salen ae Sennen) bone eee
Santos, Stuart L., Randy Day, and Stanley A. Rice. Onuphis simoni, a new
species of polychaete (Polychaeta:Onuphidae) from south Florida ____--__________
Savage, Jay M. The systematic status of Central American frogs confused with
FACULNETOAGEHY IES CrUGrTU Sj 92 we ee a a
Shelley, Rowland M. A new xystodesmid milliped genus and three new species
from piedmont South Carolina (Polydesmida: Xystodesmidae) ____________________
Shisko, John F. Five new polychaetes of the families Eunicidae and Onuphidae,
collected in 1975 and 1976 during the southern California Baseline project _____-
Sieg, Jiirgen. A new species of the genus Paratanais (Crustacea: Tanaidacea),
JP, SVCCTOWOROMS, Viton Seemnouint Wen

437-444

731-738

1163-1173

101-106

43-5]

363-373

380-390

873-877

391-403

450-454

1110-1130

503-513

67-75

404-412

1-17

1068-1075

1233-1236

761-770

1219-1225

663-668

413-420

949-967

968-983

1271-1278

Sieg, Jurgen, and Richard N. Winn. The Tanaidae (Crustacea: Tanaidacea) of
Calitomiay withia key tothe worldigenenay sas seer ae ee ee
Smith, David G., James E. Bohlke, and P. H. J. Castle. A revision of the
Nettastomatid eel genera Nettastoma and Nettenchelys (Pisces: Anguilliformes),
withidescriptions oftsix Mewsspecies (ewan ah ieee ee
Springer, Victor G. Notes on Blenniid fishes of the tribe Omobranchini, with
deSchiptionsiof tWoMnew SPECIE Siac ees a in ee Se eee
Standing, Jon D. Occurrences of shrimps (Natantia: Penaeidea and Caridea) in
Central Calitommiaiand Oregon} so... Suis hen begs oe ee
Stock, Jan H., and Glenn Longley. The generic status and distribution of
Monodella texana Maguire, the only known North American Thermosbae-
PACS evry | iese ih Heal Bs oct sam payee) Soe a ay eds en ili FPN eee ae ee
Thurman, Carl L., Il. Uca marguerita, a new species of fiddler crab (Brachyura:
Ocypodidae) Mromileastenme Me xi Coes sae ee een once
Toll, Ronald B., and Steven C. Hess. A small, mature male Architeuthis
(Cephalopoda: Oegopsida) with remarks on maturation in the family ____________
Turner, Richard L., and David B. Campbell. A redescription of Echinaster
modestus Perrier, 1881 (Asteroidea: Spinulosida) from the eastern Gulf of
Mexico) andthe WestIndies®¢ ax: su aii waved ee Uh eee 9s A
Tyler, James C., and Keiichi Matsuura. Comments on the osteology of balis-
toid fishes (Tetraodontiformes), with notes on the triodontid pelvis ______________
Vari, Richard P., and Jacques Géry. Nannocharax maculicauda, a new species
of African characoid fish (Characoidea: Distichodontidae) with comments on
thesgenus Hemigrammocharax — asa aiai ne ae einer ere eee eee
Villalobos-Figueroa, Alejandro, and Horton H. Hobbs, Jr. A new dwarf crayfish
from the Pacific versant of Mexico (Decapoda: Cambaridae) _____________________-
Voss, Gilbert L. A redescription of Octopus ornatus Gould, 1852 (Octopoda:
Cephalopoda) and the status of Callistoctopus Taki, 1964 __________________________
Ward, Linda A. Spionidae (Polychaeta: Annelida) from Hawaii, with descriptions
OF fiVE INEWUSPECIES. 2 ea et ah) eae ES eet ed eel ae
Watling, Les, and Heather Holman. Additional acanthonotozomatid, param-
phithoid, and stegocephalid Amphipoda from the Southern Ocean ________________
Weems, Robert E., Kenneth A. Beem, and Timothy A. Miller. A new species
of Bothriolepis (Placodermi: Bothriolepidae) from the Upper Devonian of Vir-
Pina CWSA) jw bear View weahs Ty snails eer A a ee
Wicksten, Mary K. The species of Automate (Caridea: Alpheidae) in the eastern
Pacific Oceani paeciess. soth.at een iiree hh” ae Aen. tee ee
Williams, Lucy Bunkley, and Ernest H. Williams, Jr. Nine new species of
Anilocra (Crustacea: Isopoda: Cymothoidae) external parasites of West Indian
coralireef fishes: 04-8220 o..2 22 Reg Eee, Soh See
Wilson, George D. Taxonomy and postmarsupial development of a dominant
deep-seaeuny copid isopod: (Crustacea) sesame ee ee ee
Young, David N. Taxonomic observations on eastern Pacific Antithamnion
species (Rhodophyta: Ceramiaceae) described by E. Y. Dawson ________-___---__-

Vili

315-343

535-560

699-707

774-786

569-578

169-180

753-760

344-351

52-66

1076-1084

492-502

525-534

713-730

181-227

984-1004

1104-1109

1005-1047

276-294

94-100

BIOLOGICAL SOCIETY OF WASHINGTON
PROCEEDINGS

1059 Meeting—16 March 1981

The 102nd Annual Meeting was called to order by President Richard
Banks at 2:00 p.m. in the Waldo Schmitt Room of the National Museum
of Natural History, Smithsonian Institution, Washington, D.C. There were
23 members present.

The minutes of the 101st Annual Meeting were approved as published in
Volume 93, part 4 of the Proceedings.

Treasurer David Pawson submitted and summarized the Treasurer’s
Report for 1980. The total funds available in 1980 were $55,047.27, and
expenses were $49,495.16. The proposed budget for 1981 estimates income
of $32,400.00, expenses of $19,450, and $12,950 available for subsidizing
publication of the Proceedings. The Finance Committee, consisting of C. W.
Hart, A. C. Cohen, and D. L. Pawson, signed the report which was
attached to these minutes. Pawson was thanked for his service to the
Society.

The Editor’s Report, in the form of Volume 93, was completed with
the publication of Part 4 on 16 January 1981. Editor Hart, who is stepping
down, was thanked by the members in attendance. Brian Kensley is the
new Editor.

Newly elected officers for the Society are: President, R. Manning; Vice
President, P. Spangler; Treasurer, L. Knapp; Secretary, M. Bogan;
Councillors: F. Bayer, I. Canet, K. Fauchald, D. Pawson, and A. Williams.

President Banks summarized the year’s activities of the Society. The
highlight was the Centennial Dinner held on 3 December 1980. Approxi-
mately 70 members and invited guests were present for a buffet dinner
followed by a talk by Paul Oesher on early members of the Society.
Bulletin No. 4 of the Society, A Centennial History 1880-1980, was dis-
tributed at this time. The history was written by John Aldrich.

Mike Sweeney was announced as the new Custodian of Publications.

After thanking the officers, council, and members for their cooperation,
Banks turned the meeting over to the new President, Ray Manning.
President Manning called for a show of appreciation for Banks, for his
services during his term, especially in coordinating the Centennial Dinner.
The meeting adjourned at 2:18 p.m.

Respectfully submitted,
Michael A. Bogan
Secretary

INDEX TO NEW TAXA

VOLUME 94

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

FORAMINIFERIDA
Falsipatellina™ 2205) 90 Ea ee ees
SHGSErING: LO ee a Ss TR IS A IS le hd
PORIFERA
Chlonarhodensis 2 SO eR ae sor Se IRR EA Ce hc
COELENTERATA
Anthozoa
Bathytelestome: Soe. eae e) Se Ml Ee LT aS oe Be ee
Pseudoplumarellavechidnaw, ss crse Vase eT i een ak)
TWO GDELUVGA GN, cis ies apie RAR eB TN ag Nig Sa cme aes NE gee
SGN PUODOGIUIN py, 20 op. su. Wag Wie ye a Ee Oe oe Le
SECU COLEL ESI O) eae a i dR ee | Ne de eM ee ae gt
TESS CV GREGG), oe 38 ae ee Sy we eI eM eel Ee le
FHOGOTGN AN). Rie ACE PAZ 2 ON eR ae ren a
PLATYHELMINTHES
Trematoda
Paraheteronchocotyles 22.05. 2 o wget Ls Pa Bs Be I rn | i
GIN GZOMECNSUS Win ke 5 BO a Ss es SER go) ee
Potamotrygonocotyle Vo See se Re ENT ies ee a
ESQ CRUSE Werth) es Seeks 0h teh aod a Rah ent ols dal ed COIN Clee lis cA MS A
SIPUNCULIDA
Aspidosiphon (Aspidosiphon)) gosnoldi 2-23
CHAETOGNATHA
Spadella legazpichessi tke. Vk 05a as ite GT AINE aay SN En EE RR
ANNELIDA
Hirudinea
Bathybdella ses Gen ein Ss etc B cea kes AG ek | ee a Te, peed ual e St Sr
SAWS CTD CM fae re A so Sh Rie ae oO Le Ra LI a
Oligochaeta
Adelodnlus:acochlearis: 222.2320. 0 cesee os. a eee ee
Homochacta: 7aptis@e: nok ee. a ee EG oe ae Dl eee
OUTST AVULUS en a a eee ae ET OI Ug UI ic a
Spirosperma™(Spiosperniva) ree eS NN eh ete eee Uae ee ee
ViGriGhGetay coz. MISE OG SD SNS O Se oe ce ale einen ere
ISTaeliSmmecs: hf) We nr ear Pee 80 Steele RS ae ee Bake eee

TRE VAG INARI Cerne ee eee Bete ene Arr meU I MUS aah Oe Ral eS Be nly oe rie an 1051

[DOSY ey ee LN A I ge LEN DR RS A Ree i OA Nak se 1051
Branchiobdellida
BOTAN ATTN OLA Ui ONC Cre see ae pe ee eg ee ae Bae 2) ee eege 677
SCORN CL ULES siutaine eh aes eM otter lh Eh wal se Le Deh le eek Ma 691
SS DUINGOG TEAMS WCDNL CIEL LLU S| Meee tains Oe Ee ieee su ee ae 849
GILOUUOLLUS Cl Capote a te tte tl Ne Sts 9 ELLY UNE WA Watt hal oa I PS 9a 853
NERV ETRSUIS TCs, ee SS Ree ee oe Se ee eee eee ae ee ee 855
SL CES CL Coen tne DCR A SD ie Et EAN Nt OA nl LE II ie sla RS RG Hs ha 856
VA CLI{CL LT UES ee na alt cp ech NES es Sm Set RG 55 ER 1 te bi ee ls Ea Se 859
Polychaeta
EXTREM EOS. CO DARLS Ls LAE EIS FT RIS IRON AIS BS OD SS, EPs tee eT EPRI ane IE 163
AGT SROCIENWS CHAM EEO EE ENE SR Oe ae eee ee eee ee 1353)
BO PEGE S elmer oh Lasse ihe a ek oN EL RRL be io eee a ena s See 1133
HOTROS DIO. a SS SI SE I Ee a aE es ne eee ee eet oe eae a 229
DDN ONDE UST LOY TNE OR i ht ER MA rs LES Sg os Sea I We 230
REMUS (RALOMUDINS IV IOLACCA: om soxt Lie an #1 mS en eh Ole NN eye re Eee ee 763
FNS [IVA OS POLO EEO [) Ul cas eae ea tate a i a a Sve i oe ee ee 717
EVGNCS COQCO BE SSe Ie 0 Leth el AAI SPUR PIE A IIE AAO ATT SEE ie hy EATERS, Sn 986
i ELLNLOSMIT OLA SES DSR SS TE IS pe as Pe a I AS I NS, Ea Ne Ses SOS 971
FADO SCOLORMC GILL ONUGUUS |g earns Me irc Eater Me a We aes Fee PBL oe ee eee 1134
OSL TIIS “TH (ap Sa ee en sare alee Nl a A a Se ORE ON aR NASR 1137
2OMCS PODUNAATTS TT EI SS ULES OT: eee eet Seer coe 719
LDIDIOTIONIIG — 5255S ERIE, SINAN SUPET wee Ni SRR Nan OE a ONE PRT arene ts et A 829
COSMIC IE ES TS SSN SE UMN TINIE OY ITS SSNPS ry see ames Wy Me Rll Ely TE 831
RNG UI RCN OL Giew ee ome ernst eo Rene MOM MMe UME ay oe ee Lo ede toni Obaumee, Le 973
MS ODRE OMAR CO MARS ME Cp ee earn a eae Re NS sek ee Be ee alae Re Se etl Hae MS
[DUTIES TOR Te TET Sik eds Saree Nee be Sak AN Aa ane 1139
TL OU CLL Cas AOE EE OS TES eS SN SS PET NLR ID A BERRA 5 SRNL 1141
SOUPS UII TIU CTU UG ee ee ere eh ton Se Ve De ee 976
SCROLLS S LEIA EE I AA SES ELE IE BI EIS SIO att sete a 979
SAU OUIY LS STS SE A Ne i A) Re A GD et eC sea re ONO gE A 663
oO GONAEKA CON Came et Ae aR Nee in eee oe oA RIEL NS tM Sales Ee Se MERU a aie 2 Se A 720
[DULG ELE SEES SSIS IS PI IO aS ON SRT, WRG ANN TE NAN SOU ys 7 ED V22
ION OSPION (MINUS PIO) (SQNGENSI 2222220 ao. ee oe ene ee ee 834
ESCUCOSCAIIDNC OMA vSQnPILIN nn ER ee 1145
EAN COS TNO MILES GLE | CLUE Se ers Sela al a EOIN FA Dale Sale nc NM Pah en 724
CANOE CIM GACSHDCNUCIILLS me aia Nl a eum ee ae NM ee al oe eee 1149
BXCT1 CLG. Sep ee A OMT gt aera Sn a, IS iP oe a2 al NOD ue pe DA 1 EE IE SNL ls 827
DENS DOU GES OC OE LE, IE I SRN RY: LINE el ret NNT PE 828
ARTHROPODA
Crustacea
ECR GUO T YE Ee ee ee ee ee ee ioe 88
EMILACAMALSBS CLULONOLIIUGIUL meats OR nh cies caihly AeA) ARG Ter hae poh oem Sees 329
BRIA CANE ELLE I LLL ft tahoe the ta Aa ere lm ee 2 ner NO § ise eh iA es shee OI 1027
CON LAU re a ria Pit soem EA 8 Katey ey 8S Rif ines or) eee ET A SORES 1018
CHE CLOG ONUUS Wye ees eho 8 tN Mere ie AP. inl aie t-onirp nro a pyre werts Pe. a EAD Re Ne ON a 1032
COUP OY ISS ga pc a A ee ey ae enol et ene pe had se lr Ge ee hs os 2 1024

OURO GET TH Bee ENGI a and I RNS Me as LE SOPs aL See Od 1030
WOlOCEN GFE bos ao! el Oe aaeatoa 0 mae Umi ea Oe 1014
MAN TIDTUSTIS! ) DE2 =k. SO a Se ee ho 1016
DGQTLULE ee 28 pee Oe no noe Santee SN AN om molt AEN Ue eahiey A 2 Se 1037
CACCIA OTSA fEUSTES ooo Be a he RS etree ene ares RN oy ahd ar oe nie a el 585
SGLEMENSIS 2 erat 2d TENE A NO ROE ah eee er ee er 372)
Calasellus: cet ieee sesen ee pe PON Ee Re rie th Ua UR tate 5 Minion aes 866
longiis- “shears wee sore bans soar ee Pie Tee roa A rie loti Nae 867
Cambarellus-prolixws 222 sowie se waar ee Sal Seen eel ae eee Pe 492
Ceératocolax mykternastes 22228: 228s we helenae ee eee ee 514
Coralanthitnas $2206 G0 je NRL ey ata ab a Nol 01 0 ae) 503
ANGE 220 BS SIC SNS, Ge US AN a ar ee a URE ea el el 507
CNRACAVOUTAE, 2285 en TEE RE oS ae 506
IDISCIAS MUSICUS (= ooo OE ea oe de 787
Eobroleus chiumashi- 5 ee Ee ee eS 301
Eudactylinaichilensis 2222.22 70) Se PE ee ee ee 745
Bury COpe ‘iphthinia: (si so0 ons Ra ee) a Ee el a 277
Gnathiphimediasexdentata, in Genta) as 193
sexdentata sexdentata 22.3. sco a ee 189 |
Gonodactylus lightbourni. «233 2. 222i Ee ee ee 708
SUQIIVCTUSUS: S20 ies <1 Vase eA pat IL Se AMM, OU alan ce 0% Beene ant Ve i 479
GOrZONISGUS, 2.8) Ne De ao Ue eh ee Eee ee le 123
LNCUSOMQGCLYLUS. oo ahead 124
Miexisquulla: 2252 LOS Tae oe Seen ene ne 595
Odimusrantarctieus, 2200 ee Ey a 205
Odontozona -libertae: 2062s UT Se Gas Co a ae 153
Oithona wellershausi. ee ee 1244
ParatanaiS Spinanotandus 2-028 202 Ee a WTA
Perichmenesb@yeri 25. 22 Sa I eS 792
Platysquilloides 00 Gay BI I SS PE OE I 593
POntOniay Viren ie. 8502 25 eh OS OA ETS hee A Oe Ee 796
Pontoniopsis paulae: 222-00 ee so 139
SULT TATU a Se ee ree ee 510
BOlUMIANONUNE So Fae NE Se ee 510
SCalpellonisS Cus c. o e a ee 1259
bimoculiscnic, 25 S26 ee a a 1264
DOCTUCULVTUES O28 oo NI oe NE Saha ND CA i 1259
Solenoceta alfonso” 2 eo ee 631
Synaptotanais nOota@bilis” co a 323
TegasteS: Acroporanus . 22.2 oe EE Re ee 254
Wicanmanewenitas a. re Soe eae gree we rk ae 171
Veleroniopsis. 2000 528 PE alah a ial wep We Ted a ae 145
Kimally ne scone ek I tas 147
Insecta
VAN TVCRON CLOG 4 ic te so so) 2 BT o egin Ne SR 1280
QVC 1 285 SES SN Te a aN a 1289
WS CUCU esata 2 Set alee TO OP Ae ancl MT AS BIN CE ele RE ee 1283
PEONUISHINa MiG) 26220 es Ee 1301
POUING Obes kee Deo 28S soe See Ss ee Same, DiS A a 1294

Xil

TLCS O11 (ite sTyLe Ss cake as ge a a he RS SD 1290

PSICUAO JCS CLG Cl pein IAA Sat a 0 lel A IN A ga ISRO 1286
POTOSI Ee Sige | eh TE TG NTE Ae BS Medan A I Rees PI New Many ae De 1298
SORGIGIDE SHS e peat Ger ok see LORE, NSE Mae sien alanis CU SL ey, BITC 1299
SQUIT CLL Thgecec meet aa while cb arte UU ids I GS bala naa Si cles ae 0 eli SRR 1292
VEO GS Vane ee OLE Sache Dosis epee ee a as Nae 0 SS OY 1296
Wlemtesm(Cleptes) wisfigaster soe sos a OES INN se lt 814
TLS EA CLILLL pn aba Pahoa A REA EN alias ee Ns, Stee Oe Jha) We eer oa 814
ONGOGIED LCS) sig site oe tee Ree Seu a) Sd Lack epg bes 816
CNCOGIE DTS) finite maaan ae apa lle ee S 816
Myriapoda
EP DOE CULICUN DD esa aN Lt AO EVAR I Y EES TE Pe SCE LON RON eR Rey OU UNE ren Ss OE er 953
COG UGS aemnren Maurin s ee ueent rR Raed an BUA IND UMN ew Jin stat detain Pa Me Nase ect Bal 956
COMV OULD) ES SEN TUNE NS SIE Nc at POEL SIERO nas a a Rea 959
UROL Sa SONAR PTO gay EE Das AUS AES CER LEA CMe LUST 961
MOLLUSCA
Gastropoda
LEP QUUDTU OUI OS ss a SAP aie REN RO SNM Dg A A cee: Mea Rr aC ee ae 439
ETRE UDI OILY cae Be SATU DO SUT Nh Ng 4 A) Se Seg NO <a Wee IGN ge 442
Harel SIN IgA CGT CLIE ame ie arc enna Nene Fa a SA yd lle ale aaah yeas eg a i 1114
[ORO GAOI se SN ER PAN aa es Scoala eta role EE OR ni De eine Dem So MS
GCUCTILEL CLD US pen Deel et esc eR AeBAINS a Sie Sieicn terme AD NW Uy Le Gy ine) a allo rym Wd ets LF
CHROME (DUTOCUAG Sp eee CS I SG ae oe a he eee ale a ne ee BL
Nodilittormal(Granulittorina) hawatiensis n.c. 22 1234
ECHINODERMATA
Asteroidea
EVO PLOSOMAESCOnDIO eek Mel ona hsn ON ns ae OVA Mg gis fe IN IN Musa he oe Voie 561
Holothuroidea
RHolothuntay(Platyperona)srowel s.-825 nies. sea ie eee tee SY a 873
HES OM GIUNAR/I LO SURG TILITT apt eltiae Sc aae Meta Dot We betes oi eee We | tg ees 450
LD ONS CLO OICT Pe rng UO Rech ea) MN de Te at ey See nN ee 397
CHES SUCIS CO! OE PEs PPS SERGE SCH) LSE TAN Se Met oA AOE SP 400
CHORDATA
Pisces
[SI EIRINAV RI DYKE] 0 & GU DYA\y SAMO ens at re Sr Mee dL Om PME DehSL wa 38
BES AIGLECURCLUGIUL DY) Sea poem OL SEUSS A REO EA Maet GN OS See Uw UMP Ie Me NRE LM te en, Sour ea 2 39
LO UEA GULTLO S Uap Ae ie en IN ok 7 HONE LUE Re ied eh EN aeten ae OE MMe gE eee oP SE em 39
ES OUNGOLSPISEVIGOINICN SIS seem ees een aU 250 Ch i ae EE le 986
@alaciwxechinamalonynchus Paes eerste a 1092
OOD SN eR AE es ds Ra Te Eke GIS GA AM ee Tee ed BIN OR AR 1096
ROIS PISS) igre eats Be ee CPA Ee NES EN Da NaS ee elit. Ae Lg Nae eye 428
NET VUCCCONUD LU Sateen catenin Winienives wenn, Ranier aA NO. LE ly Beste Se Wr oie gh eA 466
PVA CANA UST; LEP UC weet MUNA sels, Nec jek auehen EOS sy ED SVAN acyiyn Sea P yi aon ea Rie | eae 266
SOSCU OUTS ES) coe NoMa EA Se MEE EL AE RE FN Oe Ne OLA RM ae 268
ULLAL (eee Meee RE RAC RMON ee ie ADS WIN Oak AR Robe ie uel Die

Isticiobrotula olii so. Soc 2 A ee 1086

Micrognathus pygmaeus | .-....----2 4 oe ee T71
Nannocharax: moculicauda =... fs ee ee 1076
Nettastoma solitarium,..<. 15. 2.20) 22 ee ee 548
SVG OSS.» oo ESET ee bil 546
Nettenchelys exoriis : 22) cx Ve i NS ee A oh ial ae 556
BCPWY TG =o ke ee I 5)5)3)
DUOMO cs coe os See RO ION Ea OS RIS RE el 555
DY QINGCUS vio ne ee ae ene Sh
Notropis salléin.c.2. 22... be a ee 29
©mobranchus robertst 2 Sos a ee 701
OMOX' LUPUS: 2025. oo fhe ee i EN BUS ee ee Ole Se Oe Re 704
SacCOgasten vel a@NOmy ler asi a sea a hi oe ll ae oe we eh i 374
Amphibia
Dendrobates wivyersi: wo se i Se ee ene Ee Sere ee 67
Eleutherodactylus savagel: 222) a) eS ee eee 404
Aves
Chrysothly pis:chrysomelas titanoia, eee 368
Euphoniaxanthogasten oressinoma = se ee 102 ,
Schizoeaca harterti bejaranot'!2220.15.5 asd 2 ee ase ee 1068
Sporophila angolensis: ochrogyne: 2222) 8) ancy es 47
angolensis theobromae +: 2 ae ee 50
MAXIMIlAN: PATKeTeM "Ce sees NOU ee ee ol tle eae 2 45
MICROPROBLEMATICA
Pilitgella 0 OEE AS ny oa ons 4) dD an aie dea! Sa eee Te eh 421
CRYPTOGAMIA
Rhodophyta
Antithamnionella mcnabbil nic. 2-2.) Sel ee a ee oF),
Balliella pséudocorticata n.c, 22.05.22. 2-2 ee ee 94

xiv

574,06 73

Proceedings
of the
BIOLOGICAL SOCIETY
of
WASHINGTON

MAY 2.91981
LIBRARIES

Volume 94 19 May 1981 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

Associate Editors

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

Insects: Robert D. Gordon

Membership in the Society is open to anyone who wishes to join. There are no prerequisites.
Annual dues of $10.00 include subscription to the Proceedings of the Biological Society of
Washington. Correspondence concerning membership should be addressed to the Treasurer,
Biological Society of Washington, National Museum of Natural History, Smithsonian Insti-
tution, Washington, D.C. 20560. |

The Proceedings of the Biological Society of Washington (USPS 404-750) is issued quarterly
in February, May, August and November.

Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biological
Society of Washington, National Museum of Natural History, Smithsonian Institution, Wash-
ington, D.C. 20560. ;

Known office of publication: National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560.

Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044

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

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 1-17

AN HISTORICAL SYNTHESIS OF MEXICAN
MAMMALIAN TAXONOMY

José Ramirez-Pulido and Marie Claire Britton

Abstract.—The known terrestrial mammalian fauna of Mexico is com-
posed of 10 orders, 32 families, 142 genera, and 436 species. Among these
species, 160 are monotypic and 276 are polytypic comprising a total of 1033
subspecies; the total of species and subspecies is 1197. Among the 142
genera and 436 species, 6 and 142, respectively, are endemic. The type
locality of 923 categories, among the 1193 found in Mexico, lies in Mexican
territory. For the remainder, the taxa were described from specimens cap-
tured outside the country. By a historical analysis of the 923 taxa named
from Mexico, we have delineated four essential periods. The first (1831-
1881) was contemporary with the exploration of the western United States.
In this period 77 categories were described, 27 of them by Henri de Saussure
and Spencer Fullerton Baird. The second period (1887-1919) saw the awak-
ening of interest in research in Mexico by museums and governmental agen-
cies of the United States. This period ended with the Mexican Revolution
and the first World War. Four hundred and sixty categories were de-
scribed, of which Merriam classified 35%. The third period (1922-1942)
saw the renewal of exploration trips inside Mexico. One hundred and eighty-
four categories were described and it may be considered Nelson and Gold-
man’s period for they classified 52.2% of the taxa named during this period.
The fourth period (1943—present) represented a time of synthesis. Families,
genera and species complexes were revised. One hundred and ninety
categories were described of which Huey classified 28 and Goodwin 22. In
all, 73.4% of the mammalian fauna native to Mexico was described by 18
authors, and 101 other authors named the remainder. Marine mammals are
excluded from this report.

When one talks about the mammals of a certain region or country, several
questions come to mind, for example: how many generic, specific and sub-
specific categories are there, and what are they? Which of them have be-
come extinct, or are endemic, monotypic, or polytypic? What period was
the most prolific in the classification of these taxa? How many authors have
participated in the classification? Although the available information on
Mexican mammals is extensive and diverse, it has not been summarized in
order to answer the questions mentioned above.

The starting point for the preparation of this paper is the monumental
work by Hall and Kelson (1959) along with the revisions of Handley (1959),
and Van Gelder (1959a, 1959b). From that time, up to the end of 1978, the

2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

cutoff date for the incorporation of information, an extensive list of refer-
ences about the present-day state of Mexican mammalian nomenclature has
appeared. Nevertheless, we will only include the most relevant ones ac-
cording to proposed nomenclatorial changes.

Among the recent literature pertinent to the historical analysis of Mexican
mammalian nomenclature, we must mention the following works, in order
to perceive recognized categories and names. We have not made any per-
sonal taxonomic judgments, our sole intention being to gather the results of
the following authors:

Works that mention the description of new taxa: Alvarez (1962a, 1962b),
Alvarez and Ramirez-P. (1968), Anderson (1972), Baker (1967), Banks
(1967a, 1967b), Bogan (1978), Bradley and Cockrum (1968), Callahan and
Davis (1977), Carleton (1977), Davis (1969, 1970a, 1970b), Davis and Carter
(1978), Findley and Jones (1967), Gardner (1962, 1966), Genoways (1971),
Genoways and Jones (1968a, 1969b), Goodwin (1963, 1964, 1966), Hall
(1962), Hall and Alvarez (1961), Handley (1960, 1966), Huey (1960a, 1960b,
1960c, 1964), Jones (1964), Jones and Genoways (1967), Jones and Lawlor
(1965), Jones and Phillips (1964), La Val (1973a), Lee and Schmidly (1977),
Lidicker (1960b), Musser (1964), Packard (1960), Phillips and Jones (1971),
Pine (1966), Robertson and Musser (1976), Roth (1976), Russell (1968a),
Schaldach (1966), Schaldach and McLaughlin (1960), Van Gelder (1959a).

Works that propose changes in nomenclature: Alvarez (1961, 1963), An-
derson (1962, 1972), Anderson and Gaunt (1962), Anderson and Nelson
(1965), Baker (1969), Banks (1967a), Best (1978), Birney (1973), Callahan
(1977), Carleton (1977), Choate (1970), Davis (1965, 1968, 1969, 1970a,1970b,
1973), Davis and Baker (1974), Davis and Carter (1962), Diersing (1976),
Eger (1977), Findley and Jones (1967), Forman et al. (1968), Gardner (1973),
Genoways (1973), Genoways and Jones (1968b, 1969a, 1971), Goodwin
(1961, 1969), Hall (1960, 1968), Hall and Dalquest (1963), Hall and Geno-
ways (1970), Hall and Jones (1961), Hall and Ogilvie (1960), Handley (1959),
Hayward (1970), Hennings and Hoffmann (1977), Hoffmeister (1974), Hoff-
meister and de la Torre (1961), Hoffmeister and Diersing (1978), Hooper
(1972), Hooper and Musser (1964), Jones (1966), Jones and Alvarez (1962),
Jones and Carter (1976), Jones et al. (1971), Kortlucke (1973), La Val (1973a,
1973b), Lawlor (1969, 1971a, 1971b), Lee and Hoffmeister (1963), Lidicker
(1960a), Long (1972), Machado-Allison (1967), Musser (1964, 1968, 1969,
1971), Ojasti and Linares (1971), Packard (1960), Paradiso (1967), Patten
and Findley (1970), Patton and Dingman (1970), Pine (1967, 1972), Pine et
al. (1971), Russell (1968a, 1968b), Schaldach (1966), Schmidly (1972), Schmid-
ly and Hendricks (1976), Smith (1970), Van Gelder (1959b), Waithman and
Roest (1977), Wetzel (1975), Williams (1978), Wilson (1973), Yates and
Schmidly (1977), Zimmerman (1970).

In this paper, the analysis of the terrestrial mammals of Mexico is divided

VOLUME 94, NUMBER 1 3

into two parts. The first includes the whole mammalian fauna of the country,
1.e., Species and subspecies inhabiting Mexico, even though only marginally.
The second group comprises those species or subspecies whose type locality
lies in Mexico.

With the help of the Computation Services Coordination of the Univer-
sidad Autonoma Metropolitana-Iztapalapa (UAM-I), we developed a pro-
gram which enabled us to obtain a series of results among which are the
number of categories described per year, per author per year, for Mexico,
per state, and those with geographical distribution in Mexico. The orders
and families are mentioned in phylogenetic order, and the genera of each
family are in alphabetical order. Subfamilies are not mentioned.

The terrestrial mammalian fauna with geographical distribution in Mexico
is constituted of 10 orders, 32 families, 142 genera, and 436 species. Among
this diversity of species, 160 are monotypic and 276 are polytypic and in-
clude 1033 subspecies; these added to the monotypic forms make a total of
1193 (Table 1). This number could increase or decrease with the revision of
genera and families, and with a better understanding of those species and
subspecies with taxonomic problems.

Among the 142 genera in Mexico, 6 are endemic (Table 1): 1 phyllostom-
atid, Musonycteris; 1 leporid, Romerolagus; 1 geomyid, Zygogeomys; and
3 cricetids, Nelsonia, Neotomodon and Xenomys.

Among the 436 species, 142 are endemic, as follows.

One marsupial: Marmosa canescens.

Nine insectivores: Cryptotis magna, C. mexicana, Megasorex gigas, So-
rex juncensis, S. macrodon, §. milleri, S. oreopolus, S. sclateri and S.
stizodon.

Fourteen chiropterans: Musonycteris harrisoni, Artibeus hirsutus, Myotis
carteri, M. findleyi, M. fortidens, M. milleri, M. peninsularis, M. planiceps,
M. vivesi, Plecotus mexicanus, Rhogeessa alleni, R. gracilis, R. mira and
R. parvula.

Seven leporids: Romerolagus diazi, Lepus flavigularis, L. insularis, Syl-
vilagus cunicularius, S. graysoni, S. insonus and §S. mansuetus.

Eleven sciurids: Ammospermophilus insularis, Cynomys mexicanus, Eu-
tamias bulleri, Sciurus alleni, S. colliaei, S. nayaritensis, S. oculatus, Sper-
mophilus adocetus, S. annulatus, S. madrensis and §S. perotensis.

Twelve geomyids: Geomys tropicalis, Orthogeomys cuniculus, O. lanius,
Pappogeomys alcorni, P. bulleri, P. fumosus, P. gymnurus, P. merriami,
P. neglectus, P. tylorhinus, P. zinzeri and Zygogeomys trichopus.

Eighteen heteromyids: Perognathus anthonyi, P. arenarius, P. artus, P.
dalquesti, P. goldmani, P. lineatus, P. pernix, Dipodomys gravipes, D.
insularis, D. nelsoni, D. phillipsii, Heteromys gaumeri, H. goldmani, H.
lepturus, H. longicaudatus, H. nelsoni, H. temporalis and Liomys spec-
tabilis.

4 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Terrestrial mammals with geographical distribution in Mexico. The genera pre-
ceded by an asterisk are endemic to Mexico.

Number of species
Order SS Additional
Family Genera Monotypic + Polytypic = Total subspecies

Marsupialia Caluromys ~ 1 1
Didelphidae Chironectes ~ 1 1
Didelphis — 2 2

Marmosa - 2 Zz

Philander _ 1 1

Insectivora Cryptotis 4
Soricidae Megasorex ! —
Notiosorex — 1

Sorex 6 6 l

—

Scapanus - l

Chiroptera Balantiopteryx 1 l
Emballonuridae Centronycteris — 1
Diclidurus l

Peropteryx -

Rynchonycteris ] =

Saccopteryx | 1

2
3
Talpidae Scalopus - ] 2)
2
2)
1

Nw |
NK NY — — LY
N |

—
—

Noctilionidae Noctilio = 1

Mormoopidae Mormoops - ]
Pteronotus 1 3

LR —
oa

Phyllostomatidae Anoura —
Artibeus 2
Carollia 2
Centurio -
Chiroderma —
Choeroniscus 1
Choeronycteris 1 -
Chrotopterus = 1
Desmodus -
Diaemus l -
Diphylla =
Enchisthenes 1
Glossophaga 2 |
Hylonycteris —
Leptonycteris 2 —
Lonchorhina -
Macrophyllum 1 ~
Macrotus 1 1

Micronycteris 3 1

1 1
1

NO =| =|] WN —
—

Mimon
*Musonycteris

Phylloderma _ 1

Phyllostomus — 1

= | SB VN BRN KK KH NY S&S WR BR RK BS eS BS BSB NO eS BH YA)

VOLUME 94, NUMBER 1

Table 1.—Continued.

Order
Family

Natalidae
Thyropteridae

Vespertilionidae

Molossidae

Primates
Cebidae

Edentata
Myrmecophagidae

Dasypodidae

Lagomorpha
Leporidae

Rodentia
Sciuridae

Geomyidae

Genera

Sturnira
Tonatia
Trachops
Uroderma
Vampyressa
Vampyrodes
Vampyrops
Vampyrum

Natalus
Thyroptera

Anthrozous
Eptesicus
Euderma
Idionycteris
Lasionycteris
Lasiurus
Myotis
Nycticeius
Pipistrellus
Plecotus
Rhogeessa

Eumops
Molossops
Molossus
Promops
Tadarida

Alouatta
Ateles

Cyclopes
Tamandua

Dasypus
Lepus

*Romerolagus

Sylvilagus

Ammospermophilus

Cynomys
Eutamias
Glaucomys
Sciurus
Spermophilus
Tamiasciurus

Geomys

y}

—

N |

Ww |

Number of species

py
Be NOK NW BA

Oe ra

—

i)

Monotypic + Polytypic = Total

ee DO oe)

SBN Ne nH na NNN

—

eB One PRN DH MN

ie)

Additional
subspecies

3

— — YW —

he We aD

Table 1.—Continued.

Order
Family

Heteromyidae

Castoridae

Cricetidae

Erethizontidae

Dasyproctidae

Carnivora
Canidae

Ursidae

Procyonidae

Mustelidae

Genera

Orthogeomys

Pappogeomys

Thomomys
*Zyzogeomys

Perognathus
Dipodomys
Heteromys
Liomys

Castor

Baiomys
Microtus
*Nelsonia
Neotoma
*Neotomodon
Nyctomys
Ondatra
Onychomys
Oryzomys
Otonyctomys
Ototylomys
Peromyscus
Reithrodontomys
Rheomys
Scotinomys
Sigmodon
Tylomys
*Xenomys

Coendou
Erethizon

Agouti
Dasyprocta

Canis
Urocyon
Vulpes

Ursus

Bassariscus
Nasua
Potos
Procyon

Conepatus
Eira
Galictis
Lutra

Number of species

N

—

— me bo

— — = W Noe KS VY LY

Monotypic + Polytypic = Total

4
9
2
1

20

~—

—

—= N
BP NQF NHK BSP eK wenn ene np

— jp

— — LO NO —

— — — W Woe NON fB

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Additional
subspecies

20

—

VOLUME 94, NUMBER 1 7

Table 1.—Continued.

Number of species

Order SS Additional
Family Genera Monotypic + Polytypic = Total subspecies
Mephitis - 2 2 7
Mustela = l 1 9
Spilogale ~ 2 2 10
Taxidea = 1 1 1
Felidae Felis ~ 5 5 21
Lynx _ 1 1
Perissodactyla
Tapiridae Tapirus 1 _ 1 =
Artiodactyla Dicotyles — 1 1
Tayassuidae Tayassu - 1 1 1
Cervidae Mazama - 1 1 2
Odocoileus = 2 2 19
Antilocapridae Antilocapra — | l 3
Bovidae Bison — 1 1 1
Ovis - 1 1 3

Sixty-five cricetids: Nelsonia neotomodon, Neotoma alleni, N. angus-
tapalata, N. anthonyi, N. bunkeri, N. bryanti, N. goldmani, N. martinensis,
N. nelsoni, N. palatina, N. phenax, N. varia, Neotomodon alstoni, Ory-
zomys fulgens, O. melanotis, O. nelsoni, O. peninsulae, Peromyscus azte-
cus, P. banderanus, P. bullatus, P. caniceps, P. chinanteco, P. dickeyi,
P. eva, P. evides, P. furvus, P. guardia, P. hooperi, P. hylocetes, P.
interparietalis, P. lepturus, P. madrensis, P. megalops, P. mekisturus,
P. melanocarpus, P. melanophrys, P. melanotis, P. oaxacensis, P. ochra-
venter, P. pembertoni, P. perfulvus, P. polius, P. pseudocrinitus, P.
sejugis, P. simulatus, P. simulus, P. slevini, P. spicilegus, P. stephani,
P. thomasi, P. winkelmanni, P. yucatanicus, P. zarhynchus, Reithrodon-
tomys burti, R. chrysopsis, R. hirsutus, R. spectabilis, Rneomys mexicanus,
Sigomodon alleni, S. leucotis, Tylomys bullaris, Xenomys nelsoni, Microtus
oaxacensis, M. quasiater and M. umbrosus.

One dasyproctid: Dasyprocta mexicana.

One ursid: Ursus kennerleyi.

Two procyonids: Nasua nelsoni and Procyon pygmaeus.

One mustelid: Spilogale pygmaea.

Of the 1193 categories of terrestrial mammals occurring in Mexico, 878
have a precise type locality, 15 have been later restricted, 29 have only the
state or region or an even more general description and | is unknown, but
probably was also in Mexico. Assuming that all these taxa were described

8 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

500

50 400

VXVL GIIVA 40 Y3S8WAN SALLY INWND

NUMBER OF VALID TAXA

ers eee |

1758
1830 1830 40 50 60

40 50 60 70 1978

Fig. 1. Number of species and subspecies of mammals described from Mexico from 1758
to 1978. The bars represent the number of currently accepted names, and the line the cumu-
lative number of taxa.

with Mexican specimens, we have a total of 923 (Fig. 1) that represents
77.4% of the total of the Mexican terrestrial mammals.

In the remaining 270 categories, the type locality in the original descrip-
tion lies outside of Mexican territory. In 242, the type locality is precise, in
23 it is not, in 4 it was later restricted, and in | it is unknown.

An historical analysis of the 923 taxa of mammals native to Mexico, con-
sidering the order in which they were reported in the literature, permits us
to acknowledge arbitrarily 4 fundamental periods: the first from 1831 to
1881, the second from 1887 to 1919, the third from 1922 to 1942, and the
fourth from 1943 to the present (Fig. 1).

Before 1831, 12 species were described by Linnaeus, Erxleben, Gmelin,
Kerr, Cuvier, Richardson and Wagler (Table 2), based on specimens from
America which arrived at European museums through various routes. These
forms are now recognized at the subspecific level. The original information
that accompanied these specimens mentioned their provenance only very
generally, e.g., America, New Spain or Mexico. The type locality was estab-
lished later by restriction.

The first period (1831-1881) corresponds with the great expansion of ex-

VOLUME 94, NUMBER 1

Table 2.—First taxa described for Mexico.

Taxa

Felis pardalis pardalis

Nasua nasua molaris

Spermophilus mexicanus mexicanus
Spermophilus variegatus variegatus
Conepatus semistriatus conepatl
Odocoileus virginianus mexicanus
Coendou mexicanus mexicanus
Potos flavus prehensilis

Mazama americana temama
Sciurus aureogaster aureogaster
Thomomys umbrinus umbrinus
Lepus callotis callotis

Author and year

Linnaeus, 1758
(Linnaeus, 1766)
(Erxleben, 1777)
(Erxleben, 1777)
(Gmelin, 1788)
(Gmelin, 1788)
(Kerr, 1792)
(Kerr, 1792)
(Kerr, 1792)
(Cuvier, 1829)
(Richardson, 1829)
Wagler, 1830

ploration activity in the western United States. The first works that would
later have a decisive impact on research in the mammalian field appeared
at this time. The Viviparous Quadrupeds of America by Audubon and Bach-
man was published between 1845 and 1854, The Mammals of North America
by Baird in 1859, and Furbearing Animals by Coues in 1877.

The Smithsonian Institution was founded in 1846 and the United States
National Museum, where much of the material collected by the first natu-
ralists was deposited, was established in 1876. Among these first naturalists,
the most outstanding were Spencer Fullerton Baird and the military physi-
cians Elliot Coues and Edgar Alexander Mearns, known for their biological
works and for the influence they exercised in the following periods.

In this period, 77 forms were described. Outstanding were the works of
the Swiss naturalist Henri de Saussure who was in charge of a scientific
mission in Central America, whence he went to Mexico where he described
19 forms, and of Baird, the greatest naturalist at the end of the nineteenth
century and a pioneer of American vertebrate zoology, who described 8.

The other forms were described by the following authors: 7 by Peters; 6
by Lichtenstein (French naturalist who studied insects); 6 by Gray; 5 by H.
Allen; 4 by Coues; 4 by Wagner; 3 by Bennett; 2 each by Le Conte, J. A.
Allen, and Hays; and 1 each by Audubon and Bachman, Berlandier, Caton,
Dobson, Hamilton-Smith, Richardson, Tschudi, Wagler, and Waterhouse.

The second period (1887-1919) was marked by the awakening of interest
in exploration and research in Mexico by museums and governmental agen-
cies of the United States. Trained personnel and large resources were avail-
able, resulting in the formation of important collections of mammals from
different regions of the country.

The most important part of this period was the one between 1887 and

10 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1909. Later, between 1910 and 1919, the country was agitated by the Mex-
ican Revolution, a significant factor which explains the decrease in explo-
ration, and resulted in the low number of forms described at this time (Fig.
1). Another factor that contributed to this decrease in publications was the
first World War (1914-1918).

This period can be considered the one of Clinton Hart Merriam, not only
because of the number of taxa he classified, but also because of his super-
vision of the United States Bureau of Biological Survey, which at that time
included E. W. Nelson and E. A. Goldman, among others. In this period,
460 forms were described, of which Merriam classified 164, or 35%. Mer-
riam’s activity, along with that of Nelson and Goldman, resulted in 227 taxa,
the equivalent of 49.3% of the total described for this period.

Other researchers that stand out for the number of taxa they described,
are: J. A. Allen, 63; Nelson and Goldman, 63; Osgood, 42; Thomas, 27;
Elliott, 22 and Miller, 13. These authors described 394 of the 460 categories,
1.e., 85.6% of the mammals acknowledged for this period. The following
authors described the remaining 66: Mearns and Townsend, 9; Howell, 8;
J. A. Allen and Chapman and Bailey, 7; Andersen, 3; G. M. Allen,
Bangs, Hollister, Bryant, and Ward, 2; Lydekker, Menegaux, Hahn, San-
born, Grinnell and Swarth, Phillips, Jackson, H. Allen, Coues, Diaz, Rhoads,
Stowell, and Major, 1.

The third period (1922-1942) was marked by the renewal of exploration
trips inside Mexico. One hundred eighty-four forms were described, and it
could be considered the period of Nelson and Goldman who described 96
taxa, or 52.2% of the ones corresponding to this period.

A detailed analysis of the researchers that named these 184 forms gives:
Nelson and Goldman with 64; Goldman with 32; Burt with 25, and Huey with
23; Jackson with 6; Benson with 5; Goldman and Kellogg, Hall, Miller and G.
M. Allen with 4; Benson and Tillotson, Blossom, Howell, Orr, and Osgood
with 2; Anthony, Burt and Hooper, Maillard, Martinez and Villa, Sanborn,
Shamell, and Villa with 1.

The fourth period (1943—present) was a continuation of the preceding one,
but differs in that it was a time of synthesis. The variation between and
within populations was studied. Families, genera, and complexes of species
were revised with the help of other disciplines like cytogenetics and bio-
chemistry. Mammals were studied in their natural conditions and in the
laboratory. The number of institutions and persons with a professional in-
terest in mammalogy multiplied. In some institutions, several researchers
exerted influence directly by their academic production, or by the training
of highly qualified personnel.

There are some particularly noteworthy institutions, that have contributed
greatly to the study of the mammals of Mexico. We will only mention the
most important ones: The University of Kansas, University of Michigan,
San Diego Society of Natural History, Michigan State University, the

VOLUME 94, NUMBER 1 11

Table 3.—Selected authors and the number of presently recognized species and subspecies
of Mexican mammals they described. The period in which they described the forms is also
given.

Number

Author of taxa Period
C. H. Merriam 164 1887-1907
E. W. Nelson and E. A. Goldman 73 1907-1934
J. A. Allen 65 1877-1906
E. A. Goldman 64 1904-1945
L. M. Huey 51 190S5—1964
W. H. Osgood 46 1900-1945
O. Thomas Di 1892-1915
W. H. Burt 26 1932-1948
E. W. Nelson 25 1898-1912
G. G. Goodwin Map 1953-1966
D. G. Elliot D2, 1896-1905
H. de Saussure 19 1860-1861
R. H. Baker 16 1951-1967
G. S. Miller 13 1897-1914
R. J. Russell 12 1952-1968
E. T. Hooper 2 1947-1957
E. R. Hall 11 1948-1962
A. H. Howell 10 1902-1928

American Museum of Natural History, the Smithsonian Institution, Texas
A. and M. University, Universidad Nacional Autonoma de México and
Instituto Politecnico Nacional in México, and, recently, Texas Tech Uni-
versity, and the Carnegie Museum of Natural History.

In this period, 190 categories were described. A detailed analysis of the
authors gives: Huey, 28; Goodwin, 22; R. H. Baker, 16; Hooper, and Rus-
sell, 12; Anderson, and Dalquest, 9; Hall, 7; Davis, 5; Banks, and Russell
and Baker, 4; Alvarez, Genoways and Jones, Goldman, Handley, and
Setzer, 3; Benson, Hoffmeister and de la Torre, La Val, Osgood, Pack-
ard, Schaldach, and Villa-R., 2; Alvarez and Ramirez-P., Baker and Stains,
Bogan, Bradley and Cockrum, Burt, Callahan and Davis, Carleton, Dalquest
and Hall, Davis and Lukens, Findley, Findley and Jones, Gardner, Hall and
Alvarez, Hall and Dalquest, Hall and Villa, Genoways, Hoffmeister, Jackson,
Jones, Jones and Genoways, Jones and Lawlor, Jones and Phillips, Laurie,
Lawrence, Lee and Schmidly, Lidicker, Musser, Phillips and Jones, Pine,
Robertson and Musser, Roth, Schaldach and McLaughlin, and Van Gelder, 1.

A summary of the researchers who have formed the catalogue of mam-
mals of Mexico is presented in Tables 3 and 4. It is interesting to note that
18 of them described 678 forms or 73.4% of those described for Mexico
(923), and the remaining 245 were described by 101 authors, giving 119 as
the sum of authors who have proposed names.

12 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 4.—Number of authors that have contributed the description of fewer than ten species
and subspecies of Mexican mammals.

Taxa
Number of authors Per author Total
53 1 53)
20 2 40
8 3 24
>) 4 20
2 5 10
3 6 18
4 7 28
2 8 16
4 9 36
Acknowledgments

The authors would like to express their thanks to Mathematician Eduardo
Barroso, from the Coordinacion de Servicios de Computo of the UAM-I,
for developing the computational programs, and to Sydney Anderson and
Guy G. Musser of the American Museum of Natural History and Hugh H.
Genoways of the Carnegie Museum of Natural History, who read the first
draft of the manuscript.

Literature Cited

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. 1962a. A new subspecies of ground squirrel (Spermophilus spilosoma) from Tamau-

lipas, Mexico.—Univ. Kansas Publ., Mus. Nat. Hist. 14(8):121-124.

. 1962b. A new subspecies of wood rat (Neotoma) from northeastern Mexico.—Univ.

Kansas Publ., Mus. Nat. Hist. 14(11):139-143.

. 1963. The recent mammals of Tamaulipas, Mexico.—Univ. Kansas Publ., Mus. Nat.

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, and J. Ramirez-Pulido. 1968. Descripcion de una nueva subespecie de Spermophilus
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Anderson, S. 1962. Tree squirrels (Sciurus colliaei group) of western Mexico.—Amer. Mus.

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. 1972. Mammals of Chihuahua taxonomy and distribution. Bull. Amer. Mus. Nat. Hist.

148: 149-410.

, and A. S. Gaunt. 1962. A classification of the white-sided jack rabbits of Mexico.—

Amer. Mus. Novitates 2088: 1-16.

, and C. E. Nelson. 1965. A systematic revision of Macrotus (Chiroptera).—Amer.

Mus. Novitates 2212: 1-39.

Baker, R. H. 1967. A new subspecies of pallid bat (Chiroptera: Vespertilionidae) from north-
eastern Mexico.—Southwestern Nat. 12(3):329-330.

VOLUME 94, NUMBER 1 13

. 1969. Cotton rats of the Sigmodon fulviventer group.—Univ. Kansas Mus. Nat. Hist.,

Misc. Publ. 51:177—232.

Banks, R. C. 1967a. The Peromyscus guardia-interparietalis complex.—J. Mamm. 48(2):210-

218.

1967b. A new insular subspecies of spiny pocket mouse (Mammalia; Rodentia).—

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Best, T. L. 1978. Variation in kangaroo rats (genus Dipodomys) of the heermanni group in
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Birney, E. C. 1973. Systematics of three species of woodrats (genus Neotoma) in central
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Bogan, M. A. 1978. A new species of Myotis from the Islas Tres Marias, Nayarit, Mexico,
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Bradley, W. G., and E. L. Cockrum. 1968. A new subspecies of the meadow vole (Microtus
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Callahan, J. R. 1977. Diagnosis of Eutamias obscurus (Rodentia: Sciuridae)—J. Mamm.

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, and R. Davis. 1977. A new subspecies of the cliff chipmunk from coastal Sonora,

Mexico.—Southwestern Nat. 22(1):67-75.

Carleton, M. D. 1977. Interrelationships of populations of the Peromyscus boylii species group
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675: 1-47.

Choate, J. R. 1970. Systematics and zoogeography of Middle American shrews of the genus
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Davis, B. L., and R. J. Baker. 1974. Morphometrics, evolution and cytotaxonomy of mainland
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Southwestern Nat. 14(1):15-—29.

. 1970a. The large fruit bats (genus Artibeus) of Middle America, with a review of the

Artibeus jamaicensis complex.—J. Mamm. 51(1):105—122.

. 1970b. A review of the small fruit bats (genus Artibeus) of Middle America. Part IT.—

Southwestern Nat. 14(4):389-402.

—. 1973. Geographic variation in the fishing bat, Noctilio leporinus.—J. Mamm.

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, and D. C. Carter. 1962. Notes on Central American bats with description of a new

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, and . 1978. A review of the round-eared bats of the Jonatia silvicola complex,

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Forman, G. L., R. J. Baker, and J. D. Gerber. 1968. Comments on the systematic status of
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14 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

. 1966. A new subspecies of the Aztec mastiff bat, Molossus aztecus Saussure, from

southern Mexico.—Los Angeles Co. Mus., Contrib. Sci. 111:1-5S.

. 1973. The systematics of the genus Didelphis (Marsupialia: Didelphidae) in North and

Middle America.—Special Publ. Mus., Texas Tech Univ. 4:1-81.

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, and J. K. Jones, Jr. 1968a. A new mouse of the genus Nelsonia from southern Jalisco,

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, and . 1968b. (1971). Notes on spotted skunks (genus Spilogale) from western

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, and . 1969a. Taxonomic status of certain long-eared bats (genus Myotis) from

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, and . 1969b. Notes on pocket gophers from Jalisco, Mexico, with descriptions

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, and . 1971. Systematics of southern banner-tailed kangaroo rats of the Dipod-
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Goodwin, G. G. 1961. Flying squirrels (Glaucomys volans) of Middle America.—Amer. Mus.

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. 1964. A new species and a new subspecies of Peromyscus from Oaxaca, Mexico.—

Amer. Mus. Novitates 2183:1-8.

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—. 1969. Mammals from the state of Oaxaca, Mexico, in the American Museum of
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Hall, E. R. 1960. Oryzomys couesi only subspecifically different from the marsh rice rat,
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———. 1968. Variation in the blackish deer mouse, Peromyscus furvus.—An. Inst. Biol.,

Univ. Nal. Auton. Mexico 39, Ser. Zool. (1):149-154.

, and T. Alvarez. 1961. A new subspecies of pocket gopher (Heterogeomys) from

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, and W. W. Dalquest. 1963. The mammals of Veracruz.—Univ. Kansas Publ., Mus.

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, and H. H. Genoways. 1970. Taxonomy of the Neotoma albigula-group of woodrats

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, and J. K. Jones, Jr. 1961. North American yellow bats, *“‘Dasypterus”’ and a list of

the named kinds of the genus Lasiurus Gray.—Univ. Kansas Publ., Mus. Nat. Hist. 14

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,and K.R. Kelson. 1959. The mammals of North America.—The Ronald Press Co., New

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Handley, C. O., Jr. 1959. A revision of American bats of the genera Euderma and Plecotus.

Proc. U.S. Nat. Mus. 110:95—246.

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479.

VOLUME 94, NUMBER 1 15

. 1966. Description of new bats (Chiroderma and Artibeus) from Mexico.—An. Inst.

Biol., Univ. Nal. Auton. México 36(1—2):297-301.

Hayward, B. J. 1970. The natural history of the cave bat Myotis velifer.—Western New
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Hennings, D., and R. S. Hoffmann. 1977. A review of the taxonomy of the Sorex vagrans
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Hoffmeister, D. F. 1974. The taxonomic status of Perognathus penicillatus minimus Burt.—

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,and L. dela Torre. 1961. Geographic variation in the mouse Peromyscus difficilis.—J.

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, and V. E. Diersing. 1978. Review of the tassel-eared squirrels of the subgenus Otfo-
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Hooper, E. T. 1972. A synopsis of the rodent genus Scotinomys.—Occas. Papers Mus. Zool.,

Univ. Michigan 665: 1-32.

, and G. G. Musser. 1964. Notes on classification of the rodent genus Peromyscus.—

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Huey, L. M. 1960a. A new race of pocket gopher (Thomomys) from San Fernando Mission,

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1960b. Two new races of Perognathus spinatus from Baja California, Mexico.—

Trans. San Diego Soc. Nat. Hist. 12(24):409-412.

. 1960c. Comments on the pocket mouse, Perognathus fallax, with descriptions of two

new races from Baja California, Mexico.—Trans. San Diego Soc. Nat. Hist. 12(25):413-

420.

. 1964. The mammais of Baja California, Mexico.—Trans. San Diego Soc. Nat. Hist.

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Jones, J. K., Jr. 1964. A new subspecies of harvest mouse, Reithrodontomys gracilis, from

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, and T. Alvarez. 1962. Taxonomic status of the free-tailed bat, Tadarida yucatanica

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, and H. H. Genoways. 1967. A new subspecies of the free-tailed bat, Molossops

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, and T. E. Lawlor. 1965. Mammals from Isla Cozumel, Mexico, with description of

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, and G. L. Phillips. 1964. A new subspecies of the fruit-eating bat, Sturnira ludovici,

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Kortlucke, S. M. 1973. Morphological variation in the kinkajou, Potos flavus (Mammalia:
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16 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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Patten, D. R., and L. T. Findley. 1970. Observations and records of Myotis (Pizonyx) vivesi
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Phillips, C. J., and J. K. Jones, Jr., 1971. A new subspecies of the long-nosed bat, Hylonycteris
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Pine, R. H. 1966. Baeodon meyeri (Chiroptera: Vespertilionidae) a new species of bat from

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VOLUME 94, NUMBER 1 L/

, D. C. Carter, and R. K. La Val. 1971. Status of Bauerus Van Gelder and its rela-

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Departamento de Biologia, Universidad Autonoma Metropolitana-Izta-
palapa, Apartado Postal 55-535, México 13, D.F. Mexico.

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 18-36

SYSTEMATICS AND VARIATION OF THE AZTEC
SHINER, NOTROPIS SALLEI, A CYPRINID
FISH FROM CENTRAL MEXICO

Barry Chernoff and Robert Rush Miller

Abstract.—Notropis sallei (Gunther) inhabits upper tributaries of the Rio
Balsas, Rio Lerma, and Rio Panuco drainages, the Valley of Mexico, and
the small endorheic basin of the Rio Grande de Morelia. The Aztec shiner
has had a confusing taxonomic history, having been placed in no fewer than
eight genera and 16 nominal taxa. There is much variation within popula-
tions and overlap of ranges for most characters analyzed among disjunct
populations, thus supporting our conclusion that recognition of only a single
highly variable taxon, properly named Notropis sallei, is warranted. A re-
description is presented and the generic assignment of the Aztec shiner is
discussed.

The Aztec shiner, Notropis sallei (Gunther), is one of the southernmost
representatives of the American Cyprinidae. It is confined to the Rio Lerma
system, including its former Pleistocene connectives within the geological
limits of the Mesa Central of Mexico (West, 1964:Fig. 8). The species in-
habits upper tributaries of the Rio Lerma, Rio Balsas, and Rio Panuco
drainages, the Valley of Mexico, and the small endorheic basin of the Rio
Grande de Morelia (Fig. 1). All of these are or have been tributary to the
Pacific except the Rio Panuco, which flows to the Gulf of Mexico. Head-
water erosion by the Rio Panuco into the Mexican Plateau (Segerstrom,
1962) has transferred the Aztec shiner and other Lerma basin fishes to the
Atlantic Slope (Hubbs and Turner, 1939; Barbour, 1973; Barbour and Miller,
1978).

The minnow populations here referred to Notropis sallei had a confusing
taxonomic history over the last century, with assignment to no fewer than
8 genera, 7 species, and 16 nominal taxa (see below). Papers by Meek (1904),
Regan (1906-08), de Buen (1940) and Alvarez (1970) failed to resolve the
number of taxa represented by the Aztec shiner or to clarify their geograph-
ical distributions. This situation resulted from inadequate study material and
from different concepts of what constitute recognizable species and genera.
Not enough was known to permit proper evaluation of traits that were
thought to distinguish the Aztec shiner generically from its northern rela-
tives, especially within the species-rich genus Notropis. Although we still
need more information for confident generic assignments of American min-

VOLUME 94, NUMBER |! 19

BALSA/S

18°

1022 100° 98°

Fig. 1. Study locations (solid dots) of Notropis sallei from the Mesa Central of Mexico.
The solid star indicates the type locality of Ceratichthys sallaei; the open star shows Mexico
City in the Valley of Mexico; ‘*‘a’’ points to Lago de Cuitzeo and the Rio Grande de Morelia
drainage; Balsas is the Rio Balsas drainage; Lerma is the Rio Lerma drainage; and Panuco is
the Rio Panuco drainage.

nows, this study demonstrates for the first time that the Aztec shiner belongs
to a single taxon—a conclusion tentatively reached by Miller (1976). Gilbert
(1978) followed Miller (1976) but corrected the spelling from N. sallaei to
Notropis sallei, because the species was named for Mr. A. Sallé (Regan,
1906-08: vii).

The purpose of this paper is to determine the proper specific name for the
Aztec shiner, analyze the geographic variation of its disjunct populations,
and decide whether they represent more than one taxon. In addition, we
comment on the generic placement of N. sallei and provide a redescription.

20 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Methods and Materials

Most specimens were examined for 13 meristic and 15 morphometric vari-
ables, including those previously considered to be diagnostic among the
nominal taxa. Counts and measurements are as per Hubbs and Lagler
(1964). Vertebral counts include the Weberian complex as four and the
urostylar vertebra as one. All measurements were made with dial calipers
to the nearest 0.1 mm.

To correct for distributional properties of ratios, means and 95% confi-
dence intervals were computed from the arcsine transformed morphometric
ratios (Sokal and Rohlf, 1969). Accordingly, the upper and lower confidence
limits are asymmetric about the mean. The variability of several characters
was compared, independent of the mean, with the coefficient of variation
(CV). Principal components analysis (see Sneath and Sokal, 1973) was used
to assess concordant changes in morphometry; a priori groups are not
formed with this method. Principal components were computed from the
correlation matrix of standardized log-transformed morphometric variables
(variables in Table 1 plus eye diameter and standard length). The Michigan
Interactive Data Analysis System, developed by the Statistical Research
Laboratory at The University of Michigan, was used to perform numerical
and statistical analyses.

Specimens examined or recorded belong to the following institutions:
British Museum of Natural History (BMNH); Field Museum of Natural
History (FMNH); Texas Cooperative Wildlife Collection, Texas A&M Uni-
versity (TC WC); Tulane University (TU); University of Michigan Museum
of Zoology (UMMZ); and United States National Museum of Natural His-
tory (USNM).

Types of the following taxa were examined (the name of the Mexican
state follows each locality). Ceratichthys sallaei: Syntypes, BMNH
1868.3.3.1—3 (3 specimens), Rio Balsas at Cuernavaca, Puebla. Notropis
aztecus: Lectotype, USNM 45569 (1), Mexico City, Distrito Federal (DF);
Paratypes, USNM 47505 (1), USNM 125172 (1), FMNH 6604 (5), Mexico
City, DF. Notropis lermae: Holotype, USNM 50003 (1), Lake Lerma, Méx-
ico. Aztecula mexicana: Syntypes, FMNH 3606 (50), UMMZ 162462 (10),
USNM 55764 (85), San Juan del Rio, Querétaro.

Other material listed by drainages. RIo BALSAS: FMNH 4557 (2), UMMZ
66263 (1), Puebla, Puebla; UMMZ 124457 (13), Manantial at Hacienda de
Polaxtla, Puebla; UMMZ 172177 (115), Rio Tuxpan at Tuxpan, Michoacan;
UMMZ 173637 (3), E branch Rio Cuitzamola, W Toluca, Michoacan;
UMMZ 191695 (383), Rio Atepitzingo, SE Totimehuacan, Puebla. Rio LER-
MA: UMMZ 97439 (150), Rio Lerma near Lerma, México; UMMZ 124430
(2), Almoloya del Rio, México; UMMZ 172173 (260), Laguna Agua Blanca,
SW Toluca, México; UMMZ 192374 (4), Rio Lerma N Toluca, México;

VOLUME 94, NUMBER | 21

UMMZ 192378 (2), Rio Lerma N Toluca, México; UMMZ 192382 (2), trib.
Rio Lerma S Maravatio, Michoacan; TU 31872 (28), Rio Lerma W Atla-
comulco, México; UMMZ 193482 (3), Presa near Tabernillas, México;
UMMZ 193478 (S50), trib. Rio Lerma NW Toluca, México; UMMZ 201548
(3), Rio Lerma at Toluca, México; UMMZ 201550 (22), trib. Rio Lerma E
Zitacuaro, México. Rio PANUCO: UMMZ 193440 (8), Rio San Lorenzo at
Pena Blanca, Querétaro; TCWC 0035.1 (41), 3.5 km S Huimilpan, Queré-
taro; UMMZ 192368 (57), UMMZ 124322 (10), Rio Tula at Ixmiquilpan,
Hidalgo. VALLEY OF MExIco: FMNH 4528 (4), FMNH 4530 (5), UMMZ
108625 (192), Lago de Chalco, México; UMMZ 97440 (23), Lago de Tex-
coco, DF; UMMZ 97441 (2), Xochimilco, DF; UMMZ 189622 (135), UMMZ
192373 (369), UMMZ 192555 (39), Presa de Guadalupe, México. Rio
GRANDE DE MorRELIA: UMMZ 172178 (2), Presa de Cointzio, Michoacan;
UMMZ 182346 (23), Rio de Morelia, Michoacan.

Osteological comparisons were made from the following cleared and
stained (CS) and skeletal (S) UMMZ material: Agosia chrysogaster 162668-
CS (4); Algansea aphanea 192196-CS (6); A. barbata 194166-CS (2); A.
tincella 193665-CS (6); Hybopsis storeriana 150037-CS (4); Notropis amnis
157459-CS (11); N. atherinoides 147017-CS (16), 203986-S (19); N. boucardi
178579-CS (5); N. calientis 154338-CS (6); N. cerasinus 198728-S (2); N.
chihuahua 161735-CS (10), 161750-CS (2); N. chrysocephalus 203935-S (4);
N. cornutus 203936-S (10); N. emiliae 166119-CS (2); N. formosus 182402-
CS (5); N. galacturus 198735-S (2); N. hudsonius 163857-CS (2), 203803-S
(25); N. imeldae 188855-CS (2); N. lutrensis 113359-CS (6), 198730-S (2);
N. mekistocholas 197680-CS (2); N. nazas 161725-CS (8); N. ornatus
196726-CS (4); N. sallei 172173-CS (12), 172177-CS (6), 182346-CS (6),
189622-S (1), 191695-CS (6), 192373-CS (6); N. sp. (Mexico) 172218-CS (4);
N. stramineus 161911-CS (2), 203943-S (83); N. tropicus 192897-CS (6); N.
venustus 166280-CS (8); N. zonistius 157882-CS (11); Yuriria alta 179703-CS
(1).

Results and Discussion

The number of lateral-line scales has long been used to distinguish nom-
inal taxa here referred to N. sallei (Meek, 1904; Regan, 1906—08; Alvarez,
1970). Frequency distributions show that Rio Balsas and Rio Panuco indi-
viduals usually have more than 50 scales, whereas specimens from the Val-
ley of Mexico and Rio Lerma have fewer than 50 (Fig. 2). However, 26%
of the Valley of Mexico fish have more than 50 scales. Populations within
and among drainages exhibit a large degree of phenotypic overlap; this char-
acter clearly is not indicative of separate taxa. Rather, these data either
support the recognition of a single highly variable taxon (with a total range
of 26 counts) or are not useful for discriminating formerly recognized taxa.

DD, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Rio Lerma dr.
n-85

Valley of Mexico

n=-63

Rio Panuco dr.
n=57

Percent of Total

Rio Balsas dr.
n=-54

40 45 50 55 60 65
Lateral-Line Scales

Fig. 2. Variation of lateral-line scales among N. sallei from four drainages.

Large populational variation (15—20 scales) and broad overlap of ranges
are also manifest in the number of body-circumference scales within and
among drainages (Fig. 3). On the average, Rio Balsas and Rio Panuco fish
have more circumferential scales than those in the Valley of Mexico and
Rio Lerma (means are 43.7 and 44.8 vs. 38.0 and 38.8, respectively). Sep-
arating this character into its components, i.e. scales above and below the
lateral line (Table 1), reveals the same pattern. Similarly, the number of
scales around the caudal peduncle (Table 1) exhibits the greatest degree of
phenotypic overlap among the drainages for scale variables, and again in-
dicates the presence of only one taxon.

Comparison of numbers of precaudal, caudal and total vertebrae reveals
that ranges and means are not significantly different among populations
(P > .05; Table 1). Vertebral numbers are considerably less variable than
longitudinal or circumferential scale counts. For example, the range of CV’s
among drainages for several variables follows: total vertebrae 1.5—2.7; lat-

VOLUME 94, NUMBER 1 23

20
xg Rio Lerma dr.
n=85
O
= 20 Valley of Mexico
~ n-62
10
2 0
52044 Rio Panuco dr.
O
= ) so
an LO
O
20 Rio Balsas dr.
n-54
10

30 a5 AG 45 50
Body-Circumference Scales

Fig. 3. Variation of body-circumference scales among N. sallei from four drainages.

DS

eral-line scales 6.4-8.7; body-circumference scales 7.5—-10.0; and scales
around the caudal peduncle 7.1-10.5. Nonetheless, significant correlations
between vertebral and scale variables exist. Total vertebrae and caudal ver-
tebrae are correlated with lateral-line scales and body-circumference scales
(r > .185, 114 df, P < .05), and caudal vertebrae are correlated with scales
around the caudal peduncle (r = .259, 114 df, P < .01). Because precaudal
vertebrae are not correlated with scale variables, the trend of total vertebrae
reflects the variation of caudal vertebrae. This does not imply that precaudal
vertebrae are less variable than caudal vertebrae; the range of CV’s for each
are 2.8—4.4 and 3.1—4.6, respectively. This similarity is difficult to interpret
because it could also be due to problems in assigning the first caudal ver-
tebra. Concordant variation of scales and vertebrae was also noted by Stew-
art (1977) for Barbus radiatus.

Meek (1904:60) indicated that Aztecula lermae (Evermann and Golds-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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VOLUME 94, NUMBER | 25

borough) has nine dorsal rays; the other taxa he recognized have eight. This
was probably a misprint. We examined the types as well as other material
from the Rio Lerma system, and found no specimens with nine dorsal rays
(Table 1). Nine rays occurred, infrequently, only in Valley of Mexico pop-
ulations, and their mean was within the 95% confidence limits of the other
drainages.

Eye diameter and least depth of caudal peduncle were used by Meek
(1904) to distinguish between Aztecula vittata (=Notropis aztecus; see Mil-
ler, 1976) and A. Jermae. Rio Balsas N. sallei have smaller eye diameters;
the mean and 95% confidence limits do not overlap those from other drain-
ages (Fig. 4). However, the range of Rio Balsas populations is almost totally
contained within the ranges of Valley of Mexico and Rio Lerma N. sallei.
We consider the variability of this character to be extreme, especially for
the Valley of Mexico populations. These data show that eye diameters of
fish within and among drainages can vary by as much as 4% of the standard
length of the fish. In contrast, Rio Balsas populations tend to have the
largest average snout lengths (Table 1). The combination of small eye di-
ameters and larger snout lengths for Rio Balsas populations, and the con-
verse for the other drainages, coupled with similar postorbital lengths, has
resulted in rather uniform head lengths among the Aztec shiners from the
different drainage basins (Table 1). The data for eye diameter, snout length,
and least depth of caudal peduncle (Table 1) support recognition of only one
taxon; however, these data do indicate that, on the average, Rio Balsas
populations are somewhat modified from those of other drainages.

Alvarez (1970) distinguished N. sallei from N. lermae and N. aztecus by
the position achieved in projecting the postdorsal length anteriorly from the
origin of the dorsal fin. For example, if this projected length reaches as far
as, or anterior to, the nares, then N. sallei is indicated. This character is
really a complex variable dependent upon postdorsal length, eye diameter,
snout length, and postorbital head length. That is, the same result could be
achieved by changes in either the head components or the postdorsal length.
This complex, as well as other concurrent variations in morphology, is more
appropriately assessed by multivariate ordination procedures (e.g. principal
components analysis).

Principal components analysis identifies suites of characters that contrib-
ute to morphological variation. The character complexes indicated below
are those upon which former taxa have been based. The first principal com-
ponent explains 88.4% of the variance (eigenvalue = 13.3) and is related to
size; all variables are highly correlated with this axis (.79 < r < .99, P <
.01). Axes II and III are interesting because they elucidate trends in char-
acter variation independent of size. Axis II explains 3.4% of the variation
(eigenvalue = .51), is positively correlated with eye diameter (r = .54,
P <.05), and negatively correlated with maximum body width (r = —.26,

26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Rio Lerma dr.
" n-63
Valley of Mexico

n-48

Rio Panuco dr.
n=40
RioBalsas dr.

emptor n-44

40 45 50 95 60 65 70 VS
Eye Diameter

Fig. 4. Comparison of eye diameter (thousandths of SL) among N. sallei from four drain-
ages. Horizontal line indicates range; shaded portion is the mean + upper and lower 95%
confidence limits; and vertical point indicates the mean.

P < .05) and caudal peduncle width (r = —.33, P < .05). When projection
scores for individuals are plotted on the first two principal component axes
and clusters for populations within drainages outlined, a morphocline re-
sults, with fish having smaller eye diameters and wider bodies on the left,
and laterally compressed morphs with larger eyes on the right (Fig. 5; fish
from Rio Panuco and Rio Grande de Morelia drainages overlapped broadly
with clusters shown and were deleted from the figure for clarity of presen-
tation). These axes fail to segregate any population or those from a drainage
into a discrete cluster, thereby confirming the broad range of phenotypic
variability within and among populations, especially for Rio Lerma individ-
uals. The third principal component explains 2.7% of the variance (eigen-
value = .41) and is positively correlated with snout length (r = .20, P <
.05) and negatively correlated with caudal peduncle length (r = —.46, P <
.05) and postdorsal length (r = —.27, P < .05). When projection scores for
this axis are plotted against scores for the first two axes, discrete clusters
do not occur, and results similar to those in Fig. 5 are obtained.

These comparisons lead us to accept only one taxon, Notropis sallei.
Presumably diagnostic characters have been shown to be highly variable
among populations and do not delimit the former taxa. It is, therefore,
understandable why previous researchers having far less material for study
assigned several names to these minnows. Furthermore, we now resolve
the epithet to be sallei because one of us (RRM) examined the syntypes of
Ceratichthys sallaei and determined that this species is not a synonym of

VOLUME 94, NUMBER 1 ay)

Ceol
88.4%

Balsas
n=-44

Lerma
n-63

Mexico
n-48

AG Aue 3.4%

Fig. 5. Outlines of population clusters for N. sallei on the first two principal component
axes (I, II). The variance explained by each component is indicated.

Algansea tincella, as stated by Meek (1904:45), nor does it belong in the
genus Algansea, as placed by Jordan and Evermann (1896:212); see below.

The variation of N. sallei is comparable to, if not greater than, other
highly variable minnows, e.g. N. atherinoides (Bailey and Allum, 1962;
Resh et al., 1976), N. cornutus (Gilbert, 1964), Pimephales promelas (Van-
dermeer, 1966), Campostoma ornatum (Burr, 1976), and Barbus radiatus
(Stewart, 1977). However, each of these cyprinids occupies a much greater
geographic range than does N. sallei. The overall morphological plasticity
of the Aztec shiner may be a function of adaptations and responses to the
diverse and often isolated environments in which they occur (lakes, pools,
high- and low-gradient streams). For example, N. sallei from lakes, e.g.
Presa de Guadalupe in the Valley of Mexico, tends to have larger eyes and
is more laterally compressed (occupying the right hand side of Fig. 5),
whereas those from high-gradient streams of the Rio Balsas, Rio Lerma and

28 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Notropis sallei. A. 6 UMMZ 191695, 56.3 mm SL, Mexico, State of Puebla, Rio
Balsas drainage, Rio Atepitzingo SE Totimehuacan. B. 2 UMMZ 124322, 62.6 mm SL, Mex-
ico, State of Hidalgo, Rio Panuco drainage, Rio Tula at Ixmiquilpan.

Rio Panuco drainages are more terete, with smaller eyes (occupying the left
portion of Fig. 5).

Hubbs (1941) and Smith (1966) have noted that populations from fast
water were more attenuate than their quiet-water conspecifics. Similarly,
the third principal component axis is related to elongation of the caudal
peduncle (see above), and N. sallei from high-gradient streams has the
lowest scores on this axis (the characters are negatively correlated). Stewart
(1977) speculated that body attenuation could be associated with an increase
in vertebrae due to the influence of such factors as temperature. Scale and
vertebral variables were tested for correlation with the 2nd and 3rd principal
components to evaluate relationships among meristics and morphological
variation in the lateral and longitudinal planes, respectively. The second
axis 1S negatively correlated with body-circumference scales and scales
around the caudal peduncle (r < —.26, P < .01) and positively correlated
with precaudal vertebrae (r = .38, P < .01); the third axis was not corre-
lated (P > .05) with meristics. As the body (thorax and caudal peduncle)

VOLUME 94, NUMBER 1 uy)

B

Fig. 7. Notropis sallei. A. 6 UMMZ 193478, 63.9 mm SL, Mexico, State of Mexico, Rio
Lerma drainage, Rio Lerma NW Toluca. B. 6 UMMZ 192555, 52.1 mm SL, Mexico, State of
Mexico, Valley of Mexico drainage, Presa de Guadalupe.

increases in width, the number of circumferential scales increases, while
precaudal vertebrae decrease in number. The physical relationship between
circumferential scales and body width is clear; however, we cannot explain
the decrease in precaudal vertebrae. Elongation of the caudal region does
not seem to be associated with changes in longitudinal meristic elements
(i.e. scales or vertebrae).

Because we synonymize formerly recognized taxa with N. sallei, and our
data base significantly expands published ranges for various characters, a
redescription of the Aztec shiner follows.

Notropis sallei
Aztec Shiner
Figs. 6, 7

Codoma vittata (non Leuciscus vittatus DeKay) Girard, 1856:195 (original
description; type locality: Valley of Mexico; unavailable due to secondary
homonymy created when Gunther (1868:207) united Codoma with Leu-
ciscus); 1859:53 (redescription).—Gilbert, 1978:87 (synonymy).

30 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Ceratichthys sallaei Gunther, 1868:484 (original description; type locality:
Cuernavaca, Puebla).—Gilbert, 1978:77 (synonymy).

Hudsonius sallaei: Jordan, 1879:226 (synonymy, description).

Zophendum australe Jordan, 1880:300 (misidentification, =Algansea tin-
cella, based on our re-examination of types: USNM 23130 (3), USNM
23131 (1)). Jordan, 1890:288 (synonym of Algansea tincella).

Cliola sallaei: Jordan and Gilbert, 1883:164 (synonymy, description).

Cliola vittata: Jordan and Gilbert, 1883:172 (synonymy, description).

Notropis aztecus Woolman, 1894:63 (original description; type locality: ca-
nals around Mexico City).—Jordan and Evermann, 1896:258 (descrip-
tion).—Regan, 1906—08:160 (synonymy, description).—Alvarez and Na-
varro, 1957:23 (synonymy, description).—Alvarez, 1970:61 (keys).—
Miller, 1978:369 (may equal Ceratichthys sallaei).—Gilbert, 1978:28 (syn-
onymy).

Algansea sallaei: Jordan and Evermann, 1896:212 (description).—de Buen,
1940:14 (synonymy).

Aztecula mexicana Meek, 1902:81 (original description; type locality: San
Juan del Rio, Querétaro).—Meek, 1904:61 (description).—de Buen,
1940:21 (synonymy).—Alvarez, 1950:53 (keys).—Gilbert, 1978:61 (syn-
onymy).

Aztecula azteca: Meek, 1902:82 (listed, distribution).

Notropis lermae Evermann and Goldsborough, 1902:147 (original descrip-
tion; type locality: Lago de Lerma, México).—Alvarez, 1970:61 (keys).—
Gilbert, 1978:54 (synonymy).

Algansea tincella: in part: Meek, 1904:45 (included Ceratichthys sallaei and
Algansea sallaei as synonyms).

Aztecula vittata: Meek, 1904:59 (synonymy, description).—de Buen,
1940:20—21 (synonymy).—Alvarez, 1950:53 (keys).

Aztecula lermae: Meek, 1904:60 (synonymy, description).—de Buen,
1940:21 (synonymy).—Alvarez, 1950:53 (Keys).

Notropis sallaei: Regan, 1906—08:157 (synonymy, description).—Alvarez,
1970:61 (keys).—Miller, 1976:10 (synonymy).

Aztecula sallaei: Alvarez, 1950:52 (keys).

Notropis sallei: Gilbert, 1978:20 (synonymy, corrected spelling of epithet).

Diagnosis.—Distinguishable among all congeners by some combination
of: small scales, 39-64 in lateral line, with apical radii only; reduced pha-
ryngeal dentition, 0,4—4,0; short head with blunt snout; small fins; simple
intestine with 1 loop; head and body of breeding males with profuse small
tubercles that are larger on pectoral rays 2—8 (9) and weak on | and 9 or
10-14; maxilla without barbels; ventral surface of urohyal ovoid, without
lateral projections joining ventral midline to form V-shaped notch; ascending
process of angular not extending above % height of coronoid process of

VOLUME 94, NUMBER 1 3)

dentary; interneural elements extending posteriorly from Weberian appa-
ratus for only 2—4 vertebrae; juveniles and smaller adults with concentration
of melanophores on dorsal and ventral sides of caudal peduncle near base
of procurrent caudal rays. Notropis sallei differs from Algansea, Gila cras-
sicauda and Temeculina (subgenus of Gila), in having a tubular dermo-
sphenotic (platelike in others, except A. aphanea; see Barbour and Miller,
1978).

Description.—Measurements expressed in thousandths of SL, n = 212
except where indicated; means given in parentheses. Standard length to 80
mm; predorsal length 368-742 (573), dorsal fin originating over or anterior
to insertion of pelvic fins; preanal length 669-777 (723); posterior edge of
dorsal and anal fins straight; dorsal fin when depressed extends posteriorly
from vertical at 4 anal base to beyond anal base; prepelvic length 252-589
(531); postdorsal length 297—401 (353); caudal peduncle length 155-273 (204);
head length 245-317 (270); postorbital head length 117-155 (135); eye di-
ameter 37-77 (56); snout length 49-84 (64); upper jaw length 53-87 (70);
mouth often oblique and terminal, corner just reaches anterior margin of
orbit; maximum body depth 210-340 (269); body laterally compressed to
moderately terete, maximum width 92-209 (145); caudal peduncle least
depth 155-273 (204); caudal peduncle width 25-74 (47).

Dorsal rays 7 (5 counts), 8 (205), 9 (4); anal rays 6 (5), 7 (195), 8 (14);
lateral-line scales 39 (1), 40 (1), 41 (5), 42 (9), 43 (14), 44 (15), 45 (17), 46
Goya @9)n 48123), 49120), 50 C8) G2)752,(19)..53:(13), 545Q), 55.18):
56 (16), 57 (7), 58 (2), 59 (3), 60 (3), 61 (1), 62 (1), 63 (0), 64 (1); lateral line
incomplete to complete; body-circumference scales 30 (1), 31 (1), 32 (0), 33
(4), 34 (9), 35 (17), 36 (18), 37 (15), 38 (25), 39 (25), 40 (22), 41 (24), 42 (24),
43 (17), 44 (14), 45 (12), 46 (14), 47 (8), 48 (9), 49 (5), 50 (6), 51 (0), 52 (2),
53 (1), 54 (0), 55 (1), 56 (1); scales above lateral line 14 (2), 15 (7), 16 (32),
17 (63), 18 (51), 19 (40), 20 (19), 21 (13), 22 (15), 23 (4), 24 (2), 25 (0), 26 (0),
27 (0), 28 (1); scales below lateral line 13 (2), 14 (0), 15 (0), 16 (15), 17 (24),
S53) OG) 2060) 2 eG), 22.27), 23) 23). 4, (16) 25510), 26.07) a2
(2), 28 (0), 29 (2); scales around caudal peduncle 17 (6), 18 (14), 19 (44), 20
Girne ADs 225 69)e 255 (Zo)ne2an (24s 250 (dl) ZO) 27 Gi 28.1) cotal
vertebrae 34 (2), 35 (15), 36 (60), 37 (37), 38 (2); precaudal vertebrae 18 (7),
19 (52), 20 (54), 21 (3); caudal vertebrae 15 (2), 16 (40), 17 (62), 18 (12), 19
(1).

Color in preservation variable, from pallid to dark (Figs. 6, 7); melano-
phores always small, present on dorsal half of body, becoming concentrated
towards dorsum, not generally present below lateral line; scattered mela-
nophores on lips, becoming concentrated on snout, head, and circumorbital
regions; upper *% of opercle and 4 of preopercle with melanophores; pig-
ment lacking on intermandibular, gular and branchiostegal regions; predor-
sal stripe diffuse to distinct, expanded and intensified near dorsal-fin origin

ay) PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(often obscured in melanistic specimens); postdorsal stripe less well defined
than predorsal, expanded and intensified where it approaches anteriormost
base of procurrent caudal rays (a similar, but usually weaker, concentration
of melanophores lies directly opposite on ventral surface of peduncle, but
is frequently lacking; see Fig. 7); lateral stripe often diffuse anteriorly and
uniform or darker behind, extending posteriorly from opercle to end of
caudal peduncle; melanophores often form blotch at base of caudal rays,
sometimes extending onto caudal rays; a herringbone pattern of thin lines
of pigment extending dorsally from lateral stripe and terminating on caudal
peduncle (obscured in darkest individuals); melanophores scattered on dor-
sal, caudal and along outermost rays of pectoral fins in non-breeding indi-
viduals; breeding fish additionally with milky substance on anal fin (similar
to Cyprinella) and blotches of pigment along anal rays; breeding males with
pigment on distal portions of pelvic fins.

Breeding males, in life, with pinkish red on mandibles and throat; males
with small profuse tubercles over head and body; pectoral fins large and
rounded in males, short and pointed in females; head and snout rounded
and blunter in males than females.

Generic Status

Our placement of sallei in Notropis is provisional. The problem was
succinctly stated by Hubbs and Miller (1974:3), who noted: **. . . the present
chaotic state of generic evaluations among American cyprinids... .”’

Jordan and Evermann (1898:2799) placed Notropis aztecus in their sub-
genus Aztecula, a replacement for Azteca Jordan and Evermann (1896:258)
which was preoccupied in entomology. Meek (1904) elevated Aztecula to
generic level and referred four taxa to it (see above). Is sallei best referable
to Notropis, Aztecula, or perhaps to some other genus (e.g. Algansea)? A
similar problem was faced by Hubbs and Miller (1977) with respect to the
generic status of Dionda.

Studies by Cortés (1968), Snelson (1971), and Gilbert and Bailey (1972)
have eliminated characters previously considered to be diagnostic of No-
tropis (e.g. absence of maxillary barbels, more than four pharyngeal teeth
in the major row, a single-looped intestine, etc.). Gilbert (1978) concluded
that of six diagnostic characters, two remained: scales never numbering
more than 55 and usually fewer than 40, and standard length less than 100
mm. The inclusion of sallei in Notropis, thus, eliminates another charac-
teristic—low scale numbers. Furthermore, Notropis nazas Meek, with 44—
57 lateral-line scales (original counts of paratypes, FMNH 33572), also
proves to be an exception to the upper limit of 55 scales. Moreover, the
numerous scales of N. sallei and N. nazas tend to form a morphocline
within Notropis, rather than an hiatus. We conclude that the number of

VOLUME 94, NUMBER 1 38

Fig. 8. Ventral view of the urohyal bone of (a) N. lutrensis and (b) N. sallei.

scales (as stated above) is not diagnostic for Notropis, and that seemingly
only body size remains.

Comparison of the urohyal among selected American cyprinids yields
some interesting results. With the exception of N. amnis, N. boucardi, N.
imeldae, N. ornatus, and N. sallei, the urohyals of all Notropis examined
(see methods and materials) have the ventral surface with lateral projections
that extend up to or beyond the posterior margin of the dorsal aspect of the
bone, joining the ventral midline to form a V-shaped notch (Fig. 8). Notropis
sallei, N. amnis and N. ornatus have urohyals with an ovoid ventral surface
(Fig. 8); this surface is reduced laterally and less than ovoid in N. boucardi
and N. imeldae. The urohyals of these five species are phenetically similar
to non-Notropis examined (Hybopsis storeriana, Agosia chrysogaster, Yu-
riria alta, Algansea aphanea, A. barbata and A. tincella; also see Barbour

34 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

and Miller, 1978). We cannot determine, at this time, which of the states is
primitive or derived. However, it is clear that the five putative species of
Notropis cannot share the derived state with the other species of Notropis
examined. Furthermore, we decline to infer relationships of these five
species with other cyprinid genera because of the possibility that ovoid or
subovoid urohyals are primitive, and relationships based upon symplesio-
morphy should be avoided (Hennig, 1966). It is of further interest that N.
amnis, N. boucardi, N. imeldae, and N. ornatus are also debatably referred
to Notropis (see Clemmer, 1970; Miller, 1976, 1978; and Gilbert, 1978).

Although sallei is distinguished from many Notropis (including the type
species, N. atherinoides Rafinesque) by the shape of the urohyal, the small
scales, and its ‘“‘chub-like”’’ rather than ‘“‘shiner-like’’ appearance, we do not
propose that sallei be reassigned to Aztecula. Rather, we provisionally refer
sallei to Notropis because generic changes should not be made until rela-
tionships among diagnosable genera have been determined (clearly beyond
the scope of this paper); therefore, we maintain nomenclatural stability and
follow recent authors who have placed sallei in Notropis (e.g. Alvarez,
1970; Miller, 1976; and Gilbert, 1978).

Resumen

Notropis sallei (Gunther) habita los rios del este de la Mesa Central in-
cluyendo las cabeceras de las systemas siguientes: Rio Balsas, Rio Lerma,
Rio Panuco, y las cuencas cerradas del Valle de México, y el Rio Grande
de Morelia. N. sallei ha tenido una historia taxonomica confusa y ha sido
incluido en no menos de ocho géneros y diéciseis especies. Los caracteres
morphologicos varian mucho dentro de las diferentes poblaciones y se su-
perponen mucho entre poblaciones alopatricas. La variabilidad de las ca-
racteres diagnosticos de las especies identificadas con anterioridad también
se superponen. Este estudio muestra por la primera vez que estas pobla-
ciones comprenden una sola especie. Gilbert (1978) corregio el deletreo de
N. sallaei a N. sallei porque la especie era en honor del Sr. A. Sallé. La
posicion genérica de la especie es discutida y al mismo tiempo una rede-
scripcion es dada.

Acknowledgments

We are grateful to Clyde D. Barbour and Michael L. Smith for critically
reviewing the manuscript and offering helpful suggestions, Marco A. Gil
and Michael L. Smith for helping with the resumen, Edward C. Theriot for
skillful preparation of Figs. 6 and 7, and John D. McEachran (TCWOC),
Royal D. Suttkus (TU), Stanley H. Weitzman (USNM), and Loren P.
Woods (FMNH) for loan of specimens. Thanks are due J. Michael Fitzsi-
mons, John T. Greenbank, Kinji Kurawaka, Frances H. Miller, and Nancy
A. Neff for assistance in the field, and Clyde D. Barbour, Ellie Baker Koon,

VOLUME 94, NUMBER 1 35

and Michael L. Smith for their collections now deposited in UMMZ. Per-
mission to collect fishes in Mexico was kindly granted by the Direccion
General de Regiones Pesqueras. Funds for field work were generously sup-
plied by the Horace H. Rackham School of Graduate Studies (Proj. 291)
and the National Science Foundation (GB-6272x, GB-14871).

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nentales Mexicanas.—Sec. Marina, Dir. Gen. Pesca e Indust. Conex., México.

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, and L. Navarro. 1957. Los peces del Valle de México.—Sec. Marina, Dir. Gen. Pesca

e Indust. Conex., México, pp. 1-62.

Bailey, R. M., and M. O. Allum. 1962. Fishes of South Dakota.—Misc. Publ. Mus. Zool.,
Univ. Michigan 119:1—131.

Barbour, C. D. 1973. A biogeographical history of Chirostoma (Pisces: Atherinidae): A

species flock from the Mexican Plateau.—Copeia 1973:533-556.

, and R. R. Miller. 1978. A revision of the Mexican cyprinid fish genus Algansea.—

Misc. Publ. Mus. Zool., Univ. Michigan 155:1-72.

Burr, B. M. 1976. A review of the Mexican stoneroller, Campostoma ornatum Girard (Pisces:
Cyprinidae).—Trans. San Diego Soc. Nat. Hist. 18:127-144.

Clemmer, G. H. 1970. Systematics and biology of the Hybopsis amblops complex.—Unpub-
lished Ph.D. dissertation, Tulane University.

Cortés, M. T. 1968. Consideraciones sobre el genero Notropis y descripcion de una especie
nueva, procedente del Rio Atoyac, en Juchatengo, Oax., México.—An. Esc. Nac. Cien.
Biol., México 15:185-192.

de Buen, F. 1940, Lista de peces de agua dulce de México. En preparacion de su catalogo.—
Estac. Limnol., Patzcuaro, Trab. 2:1—66 [mimeo].

Evermann, B. W., and E. L. Goldsborough. 1902. A report on fishes collected in Mexico and
Central America, with notes and descriptions of five new species.—Bull. U.S. Fish
Comm. 21(1901):137-159.

Gilbert, C. R. 1964. The American cyprinid fishes of the subgenus Luxilus (genus Notropis).—
Bull. Florida State Mus. 8:95—194.

—. 1978. Type catalogue of the North American cyprinid fish genus Notropis.—Bull.

Florida State Mus., Biol. Sci. 23:1—104.

, and R. M. Bailey. 1972. Systematics and zoogeography of the American cyprinid fish

Notropis (Opsopoedus) emiliae.—Occ. Pap. Mus. Zool., Univ. Michigan 664: 1-35.

Girard, C, 1856. Researches upon the cyprinoid fishes inhabiting the fresh waters of the United

States of America, west of the Mississippi Valley, from specimens in the museum of

the Smithsonian Institution,—Proc. Acad. Nat. Sci. Philadelphia 8:165—213.

. 1859. Ichthyology of the Boundary.—U.S. and Mexican Boundary Survey 2(2):1-85.

Giinther, A. 1868. Catalogue of the fishes in the British Museum, London, 7:1—512.

Hennig, W. 1966. Phylogenetic Systematics.—Univ. Illinois Press, Urbana.

Hubbs, C. L. 1941. The relation of hydrological conditions to speciation in fishes. pp. 182-

195 In: A symposium on hydrobiology.—Univ. Wisconsin Press, Madison.

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Michigan Press, Ann Arbor.

, and R. R. Miller. 1974. Dionda erimyzonops, a new dwarf cyprinid fish inhabiting the

Gulf coastal plain of Mexico.—Occ. Pap. Mus. Zool., Univ. Michigan 671:1-17.

, and . 1977. Six distinctive cyprinid fish species referred to Dionda inhabiting

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Hist. 18:267-336.

36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

, and C. L. Turner. 1939. Studies of the fishes of the Order Cyprinodontes. XVI. A

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region. pp. 365-381 Jn: Wauer and Riskind (eds.), Transactions of the Symposium on the

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Regan, C. T. 1906-08. Pisces. Jn: Biologia Centrali-Americana, 8:1—203.

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variation in the emerald shiner Notropis atherinoides from the Ohio River: an apparent
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Bull. U.S. Geol. Surv. 1104-C:87-162.

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subgenus Pantosteus, genus Catostomus.—Misc. Publ. Mus. Zool., Univ. Michigan
129:1-132.

Sneath, P. H. A., and R. R. Sokal. 1973. Numerical Taxonomy.—W. H. Freeman and Co.,
San Francisco.

Snelson, F. F., Jr. 1971. Notropis mekistocholas, a new herbivorous cyprinid fish endemic
to the Cape Fear River basin, North Carolina.—Copeia 1971:449-462.

Sokal, R. R., and F. J. Rohlf. 1969. Biometry.—W. H. Freeman and Co., San Francisco.

Stewart, D. J. 1977. Geographic variation of Barbus radiatus Peters, a widely distributed
African cyprinid fish.—Env. Biol. Fish 1:113-125.

Vandermeer, J. H. 1966. Statistical analysis of geographic variation of the fathead minnow,
Pimephales promelas.—Copeia 1966:457—466.

West, R. C. 1964. Surface configuration and associated geology of Middle America, pp. 33—
83 In: Wauchope and West (eds.), Handbook of Middle American Indians, vol. 1.—
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Mexico.—Bull. U.S. Fish Comm. 14(1895):55—66.

School of Natural Resources and Museum of Zoology, The University of
Michigan, Ann Arbor, Michigan 48109.

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 37-42

BERNARDICHTHYS ZORRAQUINOSI, A NEW GENUS
AND SPECIES OF SALMONIFORM FISH FROM
THE LATE CRETACEOUS OF OREGON

Richard W. Huddleston

Abstract.—Field samples from the Bernard Formation (lower Cenoma-
nian) of Oregon contained a single well-preserved teleost otolith that rep-
resents a new genus and species, Bernardichthys zorraquinosi (Salmoni-
formes: Bernardichthyidae). A Kevex analysis of the otolith’s composition
revealed the presence of calcium, silica, aluminum, iron, potassium, and
magnesium. This is in contrast to the composition of Pliocene otoliths from
the Purissima formation which consisted only of calcium, probably in the
form of aragonite. This specimen constitutes the third reported occurrence
of Cenomanian age otoliths and the earliest reported occurrence of an otolith
in North America.

Samples collected by Bruce Welton from the basal Bernard formation,
Oregon, yielded a single well-preserved teleost otolith. The Bernard for-
mation, established by Dickinson and Virgrass (1965), lies unconformably
on Paleozoic, Triassic, and Jurassic strata and is overlain unconformably by
Tertiary volcanics. The formation is approximately 1500 feet (460 m) thick
and consists of a pebbly sandstone to conglomerate with minor amounts of
clayey sandstones and mudstones. From invertebrate data it is considered
lower Cenomanian (lower-most upper Cretaceous age) by Dickinson and
Vigrass (1965).

Although Koken (1891) first described Cretaceous otoliths from Europe,
their occurrence in North America was not mentioned until some 64 years
later when Tychsen and Vorhis (1955) reported, but did not describe, oto-
liths from the Fox Hills Sandstone (Maastrichtian age) of South Dakota.
The first description of taxa based on Cretaceous age otoliths from North
America followed ten years later when Frizzell (1965a, 1965b) described
new genera and species of the families Albulidae and Vorhisiidae, the latter
subsequently found to be a junior synonym of the family Ariidae (J. E.
Fitch, personal communication). Discussions pertaining to Cenomanian age
otoliths are restricted to reports concerning a single otolith and an otolith
impression (Gowda, 1967a, 1967b; Stinton, 1973).

The otolith terminology follows that of Frizzell and Dante (1965); see Fig.
1. Additional terms used but not shown in Fig. 1 are: ventral furrow, a
narrow depression below the sulcus parallel to the ventral margin; excisura,

38 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

DORSAL

POSTERIOR ANTERIOR

VENTRAL

Fig. 1. Diagram of sagitta of Antigonia eos (Beryciformes: Antigoniidae) showing termi-
nology used in this paper. AR, antirostrum; C, cauda; CI, crista inferior; CS, crista superior;
DA, dorsal area; O, ostium; R, rostrum; S, sulcus; VA, ventral area.

}

a notch or indentation between the rostrum and antirostrum; and collum,
a small ridge or division line between the ostium and cauda. The term otolith
is a general term for three pairs of discrete elements occurring within the
auditory labyrinth of fishes. These elements are the sagitta, asteriscus and
lapillus.

Order Salmoniformes (sensu Greenwood et al., 1966)
Bernardichthyidae, new family

Type-genus.—Bernardichthys, new genus.

The sagitta of the Bernardichthyidae differs from that of other families in
the following combination of characters: nearly straight, horizontal cauda,
lower margin of cauda straight, upper margin tapering posteriorly; short
anteriorly expanding ostium; possessing a reduced rostrum, no antirostrum;
lacking any form of depression on the dorsal area; lacking ventral furrow;
possessing a deep ventral margin; lacking well defined crista superior and
crista inferior as preserved (the sagitta displays some attrition which may
have affected the crista superior and crista inferior); sulcus situated above

VOLUME 94, NUMBER 1 39

Fig. 2. Bernardichthys zorraquinosi, n. gen., n. sp. Holotype, LACM 4483/118693, left
Sagitta.

es

horizontal midline of sagitta; all margins smooth as preserved. Additional
characters are as defined for the genus.

Bernardichthys, new genus

Type-species.—Bernardichthys zorraquinosi, new species.

Diagnosis.—Sagitta semi-ovate; dorsal margin angular; ventral margin
broadly rounded, ventral margin asymmetrical, anteroventrally skewed;
dorsal-ventral height greater than anterior-posterior length; sulcus horizon-
tal, situated nearer dorsal margin; sulcus nearly straight; cauda narrow;
dorsal rim of cauda slightly convex tapering posteriorly; ostium short, wider
than cauda; excisura absent as preserved; ostium expanding anteriorly; ros-
trum weakly developed; crista superior and crista inferior weakly developed
or absent as preserved; dorsal and ventral areas without depressions; ventral
furrow absent.

Bernardichthys zorraquinosi, new species
Fig. 2

Holotype.—Complete left sagitta, Los Angeles County Museum of Nat-
ural History (LACM) 118693; Fig. 2.

Type locality. —LACM 4483, Bernard Formation, Crook County, Oregon;
Lat. 44°06'52”N, Long. 119°40'42”W; in the NE 4 of the SE % of the NE
% of Sec. 11, T 17S, R 25E, Dayville map, 30 min. series. The site is ap-

40 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

proximately 34% mi N of Saplee, Oregon and 2 mi NE of the Bernard Ranch,
on the north side of Beaver Creek, on the southeast side of a northeast—south-
west trending low ridge, in concretionary sandstone beds outcropping on
the slopes of the ridge.

Description.—Small left sagitta, 3.76 mm antero-posterior length, 4.00
mm dorsal-ventral height, greatest thickness 0.84 mm; sagitta semi-ovate;
inner face slightly convex; greatest thickness near sagitta midpoint; dorsal
margin (see Fig. | for terminology) angular, posterodorsally inclined at a
steep angle from the mid-ostium level, reaching greatest height at mid-sag-
itta, then posteroventrally inclined at a low angle; posterior margin pos-
teroventrally inclined at a steep angle above sulcus, rounded at sulcus,
anteroventrally inclined at a steep angle below sulcus; ventral margin deep,
asymmetrically rounded, reaching greatest depth anterior of vertical midline
of sagitta; ventral margin abruptly rounded at deepest point; anterior margin
anterodorsally inclined below level of sulcus; anteriormost portion of sagitta
missing, reconstruction of this area suggesting only a weakly developed
rostrum; antirostrum and excisura not present as preserved; sulcus situated
on dorsal half of sagitta; sulcus nearly straight, horizontal; ostium and cauda
distinct; ostium short, elliptical, anteriorly expanding, open anteriorly; col-
lum represented by a low posterodorsal-anteroventrally inclined shelf; cau-
da narrow, shallow, widest near collum, ventral margin of cauda straight,
dorsal margin slightly curved, tapering posteriorly; cauda closed posteriorly,
deepest along dorsal edge, uniformly shallowing to ventral edge; crista su-
perior weakly developed as preserved; crista inferior absent as preserved;
dorsal and ventral areas smooth, lacking depressions; ventral furrow absent;
all margins smooth as preserved; outer face smooth, unsculptured.

Comparison.—The sagitta of Bernardichthys zorraquinosi differs from
other otoliths by the following combination of characters: sagitta semi-
ovate; angular dorsal margin; anterior margin with short pointed rostrum;
posterior margin sharply rounded; ventral margin deeply rounded, deepest
nearest anterior end; sulcus nearly straight, approximately horizontal, lo-
cated on dorsal half of sagitta; ostium short, open anteriorly; weakly de-
veloped crista superior; absence of depressions on dorsal and ventral areas.

The slightly elevated position of the sulcus, the small distinct ostium, and
the lack of an outward projecting flange on the dorsal portion of the sagitta
prevent association of this form with the Osteoglossomorpha. The height-
length ratio, well developed sagitta, nearly straight sulcus, and deeply
rounded ventral margin of the sagitta prevent association of Bernardichthys
with the Ostariophysi. The nearly straight sulcus, small ostium, relatively
flat outer face, thin sagitta, deeply rounded ventral margin, and presence of
a rostrum are within the acceptable limits of the Salmoniformes (sensu
Greenwood et al., 1966). Several of these characters are reflected in mem-

VOLUME 94, NUMBER 1 4]

bers of the families Osmeridae (Suborder Salmonoidei) and Argentinidae
(Suborder Argentinoidei).

Sagittae of the extant Osmeridae and Argentinidae are characterized by
a generally subangular external shape; strongly developed rostrum; straight,
horizontal sulcus; deep subangular ventral margin; ventral margin usually
reaching greatest depth nearest anterior end in the Argentinidae, nearest the
posterior end in the Osmeridae; cauda either closed or open posteriorly;
dorsal area with prominent depression in the Osmeridae, not pronounced
or absent in the Argentinidae.

Greenwood et al. (1966) defined the Salmoniformes as containing eight
suborders (Salmonoidei, Argentinoidei, Galaxoidei, Esocoidei, Stomia-
toidei, Alepocephaloidei, Bathylanconoidei, and Myctophoidei). The
straight horizontal sulcus, elevated position of the sulcus from the horizontal
midline of sagitta, lack of a strong rostrum, and ovate sagitta differentiates
the Bernardichthyidae from the Alepocephaloidei, Bathylaconoidei, Eso-
coidei and Galaxoidei. Bernardichthys differs from the Salmonoidei and
Argentinoidei primarily by the lack of a well developed rostrum, absence
of a subangular ventral margin, and the lack of shallow depressions on the
dorsal area. The narrow cauda, wider ovate ostium, and curved dorsal rim
of the cauda differentiates Bernardichthyidae from the Stomiatoidei. Among
the Myctophoidei the characters are similar to the Harpadontidae, having
the sulcus elevated to the dorsal half of sagitta, a well defined cauda margin,
and a shorter rostrum.

The Bernardichthyidae is sufficiently distinctive to prevent assignment to
any of the extant suborders of the Salmoniformes. It is advisable to leave
the subordinal position of the Bernardichthyidae as incertae sedis.

Discussion.—A Kevex analysis of the sagitta of Bernardichthys zorra-
quinosi indicated the presence of the following elements, in order of signif-
icance: calcium, silica, aluminum, potassium, iron, and magnesium. With
the exception of calcium these probably represent replacement of the orig-
inal components which have been leached out. Similar analysis of otoliths
from the Purissima formation (lower Pliocene) revealed only calcium, prob-
ably in the form of aragonite (Huddleston, unpublished data).

Etymology.—This species is named in honor of Joseph A. Zorraquinos of
Rosemead, California, for his life-long pursuit of knowledge and science.

Acknowledgments

I thank Bruce J. Welton, Los Angeles County Museum of Natural His-
tory, for bringing the otolith to my attention and making it available for
study; Joann Welton, Chevron Oil Field Research Company, for taking the
S.E.M. photograph of the otolith; Drew Haman, Chevron Oil Field Re-

42 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

search Company, for reviewing the manuscript and for his helpful comments
and suggestions; and to Chevron Oil Field Research Company for publica-
tion permission, and defrayment of publication charges. I am indebted to
John E. Fitch for his encouragement and invaluable advice throughout my
study of fossil otoliths.

Literature Cited

Dickinson, W. R., and L. W. Virgrass. 1965. Geology of the Suplee-lzee area, Crook, Grant
and Harney counties, Oregon.—Bull. Oregon Geol. and Min. Indust. 58, 39 pp.
Frizzell, D. L. 1965a. Otolith-based genera and lineages of fossil bonefishes (Clupeiformes,

Albulidae).—Senck. Leth. 46a:85-110.

. 1965b. Otoliths of new fish (Vorhisia vulpes, N. gen., N. sp. Siluroidei?) from Upper

Cretaceous of South Dakota.—Copeia 1965(2):178-181.

, and J. Dante. 1965. Otoliths of some early Cenozoic fishes of the Gulf Coast.—Jour.
- Paleontol. 39:687—718.

Greenwood, P. H., D. E. Rosen, S. H. Weitzman, and G. S. Myers. 1966. Phyletic studies
of teleostean fishes, with a provisional classification of living forms.—Bull. Amer. Mus.
Nat. Hist. 131:339-456.

Gowda, S. S. 1967a. The first fossil otolith from India.—Bull. Geol. Soc. India 4:15—17.

. 1967b. On a new fossil fish known from an otolith from the South Indian Cenoma-

nian.—Jour. Geol. Soc. India 8:119-129.

Koken, E. 1891. Otolithes. Jn Bohm, J., De Kreidebildungen des Furbergs und Sulzbergs bei
siegsdorf in Oberbayern.—Palaeontographica 38:37—40.

Stinton, F. C. 1973. Fish otoliths from the English Cretaceous.—Palaeontol. Soc. Monogr.
2:57-126, pls. 4-8.

Tychsen, P. C., and R. C. Vorhis. 1955. Reconnaissance of geology and ground water in the
lower Grand River Valley, South Dakota.—U.S. Geol. Surv., Water-Supply Pap. 1298.

33 pp.

Chevron Oil Field Research Company, P.O. Box 446, La Habra, Califor-
nia 90631 and Section of Vertebrate Paleontology, Los Angeles County
Museum of Natural History, 900 Exposition Blvd., Los Angeles, California
90007.

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 43-51

A REVISION OF THE SUBSPECIES OF SPOROPHILA
(°ORYZOBORUS”’) ANGOLENSIS
(AVES: EMBERIZINAE)

Storrs L. Olson

Abstract.—Because of similarities in plumage and morphology, and the
occurrence of several ‘‘intergeneric’’ hybrids, the genus Oryzoborus is
merged with Sporophila. Sporophila maximiliani parkesi nom. nov. is pro-
posed to replace the preoccupied name Oryzoborus crassirostris magniros-
tris Phelps and Phelps 1950. Within Sporophila angolensis there are two
groups, distinguished by the adult male plumage. In the funerea subspecies
group, currently considered monotypic, four subspecies are now recog-
nized, based on female and subadult plumages: S. a. funerea (Sclater), S.
a. salvini (Ridgway), S. a. ochrogyne new subspecies, and S. a. aethiops
(Sclater). In the angolensis subspecies group, S. a. angolensis (Linnaeus)
is the only taxon recognized east of the Andes, with S. a. torrida (Gmelin)
a synonym; S. a. theobromae new subspecies is named for a population
isolated in the upper Magdalena Valley of Colombia.

Currently there are three species admitted in the genus Oryzoborus, a
group of heavy-billed Neotropical finches in the subfamily Emberizinae.
The smallest species, O. angolensis, the type of the genus, occurs from
Mexico to Argentina and includes the nominal species O. funereus of earlier
authors. The other two species, O. crassirostris and O. maximiliani, are
considerably larger forms confined to South America, except for small dis-
junct populations of the latter species in Panama and Nicaragua. The seven
subspecies of O. maximiliani were considered to be races of O. crassirostris
until Meyer de Schauensee (1970) showed these two taxa to be sympatric,
or nearly so, over a wide range.

Despite the obvious resemblance of Oryzoborus to members of the large
genus Sporophila, in many works (e.g. Sharpe, 1888; Ridgway, 1901; Hell-
mayr, 1938) these two genera were separated by varying numbers of seem-
ingly less similar genera, although in later treatments Oryzoborus has been
placed immediately following Sporophila (Meyer de Schauensee, 1966;
Paynter, 1970).

With the exception of the angolensis subspecies group, the males of
Oryzoborus are all black in color, apart from a white speculum and white
under the wing in some taxa, so that they resemble, for example, Sporophila
americana corvina. Males of the angolensis group have chestnut on the

44 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

lower breast and belly, and males of several species of Sporophila likewise
have chestnut in the underparts. As in Sporophila, the females of Oryzo-
borus differ in plumage from males and are generally dull brownish, similar
to females of Sporophila americana and other species.

In reading through the generic descriptions given by Ridgway (1901:564,
603), which are not written in a comparative manner so that it is not evident
which characters are ‘‘diagnostic,’’ about the only difference one can find
between Oryzoborus and Sporophila is in the “‘enormously thick and
broad’ bill of the former. This is not necessarily of generic value, however.
In the proportions of the bill the differences between O. angolensis and
Sporophila americana are scarcely any greater than between O. crassiros-
tris and O. angolensis (Fig. 1). The species of Sporophila tend to have the
culmen more curved than in Oryzoborus, but there is so much variation in
bill shape within Sporophila itself (e.g. the peculiar §. falcirostris) that this
appears to be of minor importance.

Perhaps the most telling evidence for a very close relationship between
Oryzoborus and Sporophila was presented by Sick (1963:166), who found
that hybridization between species of these two genera was not uncommon
in Brazil. He discussed five wild-taken apparent hybrids between O. a.
angolensis and S. c. caerulescens, two between O. a. angolensis and S.
lineola, and one between O. a. angolensis and S. ardesiaca. Sick likewise
called attention to the fact that the differences between the smaller forms
of Oryzoborus and the larger forms of Sporophila were not nearly as great
as had previously been supposed.

The structure of the skull in Oryzoborus and Sporophila is similar, al-
though that of Oryzoborus is more robust, with a more sharply angled ros-
trum and broader anterior portions of the palatines. Again, there are striking
differences between O. angolensis and O. crassirostris. The differences in
skull structure between Oryzoborus and Sporophila are considerably less
than those between the various species of the emberizine genus Geospiza,
as illustrated by Bowman (1961).

Considering the broader generic limits now being admitted in the Passer-
iformes, I find it strange that Oryzoborus has continued to be recognized.
Although the three species of Oryzoborus appear to be more closely related
to each other than any one of them is to a particular species of Sporophila,
they are surely part of a monophyletic assemblage that includes Sporophila,
in which they represent only the extremes in size and crushing adaptations
of the bill. I believe that it is best to emphasize the close relationship of
these birds, and the fact that they constitute a single radiation, by uniting
Oryzoborus Cabanis 1851 with Sporophila Cabanis 1844. The three species
presently recognized in Oryzoborus therefore become Sporophila angolen-
sis (Linnaeus), Sporophila crassirostris (Gmelin), and Sporophila maximi-
liani (Cabanis).

VOLUME 94, NUMBER 1 45

Fortunately, this creates no problems of homonymy, as I have found no
duplication in Oryzoborus of any of the trivial names currently applied to
Sporophila. However, Kenneth C. Parkes (in litt.) has drawn my attention
to the fact that Oryzoborus crassirostris magnirostris Phelps and Phelps
1950, is preoccupied by Coccoborus magnirostris Swainson 1837, proposed
as anew name for Loxia angolensis Linnaeus 1766. In accordance with the
systematic conclusions outlined above, I propose:

Sporophila maximiliani parkesi, nom. nov.
to replace Oryzoborus crassirostris magnirostris Phelps and Phelps 1950.

Revision of the Subspecies of Sporophila angolensis

The Lesser or Thick-billed Seed Finch, Sporophila angolensis, includes
two groups that were previously regarded as separate species. In the an-
golensis subspecies group the adult males have the lower breast and belly
chestnut, whereas in the funerea subspecies group the chestnut is replaced
with black. The S. angolensis group is found in most of tropical South
America east of the Andes and in the upper Magdalena Valley of Colombia.
The S. funerea group extends from southern Mexico through Middle Amer-
ica to western Colombia and Ecuador and east to the Magdalena Valley. I
have dealt with the interesting interaction between these two groups in the
Magdalena Valley in another paper (Olson, in press). Here I shall review
the geographic variation within each of these two subspecies groups.

The S. a. funerea Subspecies Group

Because the adult males are almost entirely black, they are not useful for
showing plumage variation, and at present S. a. funerea is not subdivided.
Females and subadult males, however, show considerable geographic vari-
ation which has been overlooked until now. At one time, Hellmayr (1911)
admitted a southern race, aethiops (Sclater) based on coloration of females
and alleged differences in bill size, for birds from western Colombia and
western Ecuador, but he later considered that its recognition was not war-
ranted (Hellmayr, 1938). Doubtless contributing to past difficulties in sub-
dividing the funerea group is the fact that subadult males are more intensely
colored than females in the same population, so that comparison must be
made sex for sex; otherwise there would appear to be considerable overlap
in characters. Subadult males were not infrequently missexed by earlier
collectors, who may have made their determinations by plumage rather than
by gonads. Individuals that are apparently missexed are not difficult to de-
tect, given sufficient material. In any case, adequate series of correctly
sexed specimens permit the identification of no less than four subspecies
within the funerea group. Names are available for all but one of these,
which, as it turns out, is the most widespread form and the best represented

46 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Bills of selected species of Sporophila to show the gradation in size between typical
Sporophila and the forms of *‘Oryzoborus’’: (left to right) S. americana corvina, S. angolensis
ochrogyne, §. crassirostris, S. maximiliani nuttingi.

in collections. For the funerea group, the following descriptions, and the
lists of specimens examined, refer to females and subadult males only.

Sporophila angolensis funerea (Sclater)

Oryzoborus funereus Sclater 1859, Proc. Zool. Soc. London 1859:378. Su-
chapam, Oaxaca, Mexico.

Characters.—Females and subadult males with underparts lighter, more
brownish, and less intensely reddish than in salvini or aethiops, but darker
than in ochrogyne. Dorsum less reddish than in salvini or aethiops but not
as pale and grayish as in ochrogyne.

Range.—The Caribbean slope of southeastern Mexico, Guatemala, Be-
lize, and Honduras.

Specimens examined.—MEXICO. TABASCO: Teapa (1, USNM; 1, FM).
VERA CRUZ: La Gloria, 45 mi S of Acayucan (1, WEVZ). GUATEMALA.
Secanquim (1, AMNH); Finca Chama (1, AMNH). BELIZE (BRITISH
HONDURAS). Near Manatee Lagoon (5, CM); All Pines (2, CM); Sittee
River, Freetown (1, CM). HONDURAS. Lake Yojoa (1, CM); Planes (2,
CM); La Ceiba (1, CM).

Remarks.—The two females from Mexico are somewhat more reddish
dorsally than the remainder of the series, but the material is insufficient to
determine if this character is constant.

Sporophila angolensis salvini (Ridgway)

Oryzoborus salvini Ridgway 1884, Proc. U.S. Nat. Mus. 6:401. Los Sabalos,
Nicaragua.

VOLUME 94, NUMBER 1 47

Characters.—Females and subadult males much more richly colored than
any of the other subspecies; underparts, particularly of the subadult males,
deep reddish chestnut; most similar to aethiops but not as sooty below, and
dorsum more reddish.

Range.—Southern Nicaragua, Costa Rica except in the extreme south-
west, and the Caribbean coast of Panama east at least to western Veraguas.

Specimens examined.—NICARAGUA. Escondido River (6, USNM);
Greytown (1, USNM); Los Sabalos (2, USNM—cotypes; 1, AMNH). COS-
TA RICA. SARAPIQUI: 1.5 mi SE of Puerto Viejo (1, WFVZ); LIMON:
Limon (5, AMNH; 2, FM); Cariari (3, WFVZ); CARTAGO: Juan Vinas (1,
MCZ); PUNTARENAS: Boruca (2, MCZ). PANAMA. BOCAS DEL
TORO: Almirante (5, USNM); Changuinola (1, USNM); VERAGUAS:
Guabal, Rio Calovévora (1, AMNH).

Remarks.—The two specimens from Boruca, Costa Rica, and that from
Changuinola, Panama, while nearest salvini, appear to show signs of inter-
gradation with the following subspecies. The birds from the western Carib-
bean coast of Panama are instantly separable from those of the rest of the
country by their much richer coloration. The subspecies salvini and och-
rogyne exhibit a pattern of distribution common to a number of other
species, whereby a northern, usually darker, form extends into Panama only
along the western Caribbean coast and is replaced elsewhere in the country
by a different subspecies that may extend into southwestern Costa Rica.

Sporophila angolensis ochrogyne, new subspecies

Holotype.—USNM 410128, female, collected 25 March 1949 at Utive, Rio
Cabobré, Panama Province, Panama, by Alexander Wetmore and W. B.
Perrygo (original number 14402).

Characters.—Females and subadult males paler than any of the other
subspecies of the funerea group, underparts deep buffy, not chestnut;
brown of breast and upperparts suffused with a grayish olivaceous wash,
not reddish, and much lighter than in other forms.

Range.—Southwestern Costa Rica, Panama except the western Carib-
bean coast, and western Colombia south at least to Buenaventura and east
as far as the Magdalena River Valley.

Etymology.—Greek, ochros, pale, and gyne, female, in reference to the
pallid coloration of the females of this subspecies. The name is a noun in
apposition.

Specimens examined.—COSTA RICA. PUNTARENAS: Helechales (1,
WFVZ); 13 km S of Palmar Sur (1, WFVZ). PANAMA. VERAGUAS:
Wilcox Camp, San Lorenzo River [Caribbean slope] (1, AMNH); Sona (2,
USNM); Isla Cébaco (1, USNM); Isla Coiba (3, USNM; 1, AMNH); Pa-
racoté (1, CM); CANAL ZONE: Gatun (7, USNM); Bohio (1, USNM);

48 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Curundu (1, USNM); Juan Mina (2, USNM); Pac, K-6 road (1, USNM).
PANAMA PROVINCE: Chico (1, USNM); Utivé, Rio Cabobré (1,
USNM—holotype); ARCHIPIELAGO DE LAS PERLAS: Isla San José
(5, USNM); COLON: Colon (1, FM); Portobello (1, USNM); SAN BLAS:
Mandinga (1, USNM). COLOMBIA. CHOCO: Atrato River (1, AMNH);
Andagoya (5, CM); El Tambo (1, CM); ANTIOQUIA: Arenosas (1,
AMNH); La Bodega, N side of Rio Negrito, Highway Sonson-Narino (1,
USNM); El Pescado, 12 km below Puerto Valdivia (1, USNM); Valdivia,
Sevilla (1, USNM); Hacienda Belén, 8 mi W of Segovia (1, USNM); CAL-
DAS: Hacienda Sofia, Rio Samana (1, USNM); VALLE: Buenaventura (1,
AMNH); CORDOBA: Nazaret, 12 mi NW of Tierra Alta, Rio Sinu (1,
USNM); Tierra Alta, Rio Sinu (1, USNM); Quebrada Salvajin, Rio Esme-
ralda, Upper Rio Sinu (2, USNM); BOLIVAR: Norosi (2, USNM); Rege-
neracion, Quebrada San Marcos, Lower Rio Cauca (2, USNM); La Raya,
Rio Cauca (1, USNM).

Remarks.—Wetmore (1957) commented on the larger size of the bill in
four specimens from Isla Coiba, Panama, and felt that these birds probably
should be recognized as a distinct subspecies, an action he failed to take in
the absence of a more adequate sample. He noted one individual from the
Canal Zone and one from Colombia that also had larger bills than normal.
I examined a female specimen of S. a. angolensis from the upper Caura
River, Venezuela (CM 32398), in which the bill was grotesquely enlarged,
being almost the size of that of S. c. crassirostris, although the wing was
of normal size for angolensis. While such specimens as these represent
aberrations, the Coiba birds probably do not, but, like Wetmore, I would
prefer to see more material before naming this population.

I have been unable to distinguish females and subadult males of ochro-
gyne from those of angolensis (including torrida), although the adult males
differ markedly. Virtually the only place where this becomes a problem is
in the Magdalena Valley, where both types of adult males occur, along with
intergrades (see Olson, in press).

Sporophila angolensis aethiops (Sclater)

Oryzoborus aethiops Sclater 1860, Proc. Zool. Soc. London 1860:88. Na-
negal, Ecuador.

Characters.—Females and subadult males darker than funerea and much
darker than ochrogyne; similar to the widely disjunct salvini but more
sooty, less intensely reddish above and below.

Range.—Southwestern Colombia in Narino, and western Ecuador.

Specimens examined. —COLOMBIA. NARINO: La Guayacana (2, LSU;
2, WEVZ); Barbacoas (1, AMNH). ECUADOR. ESMERALDAS: Esme-
raldas (1, AMNH); 10 km N of Quininde on Quininde-Esmeraldas highway

VOLUME 94, NUMBER | 49

(1, USNM); LOS RIOS: Hacienda Puerto Nuevo, Abras de Mantequilla,
ca. 3 km NE of Vinces (1, USNM); GUAYAS: Bucay (1, ANSP); San
Rafael, near Tenguel, 7 km S of Balao (1, USNM); Naranjo (1, AMNH);
DEL ORO: Zaruma (1, AMNH); Rio Pindo (1, AMNH); Portovelo (1,
AMNH).

The S. a. angolensis Subspecies Group

In this group the adult males have chestnut rather than black bellies. I do
not recognize any subspecies, other than the nominate one, in the extensive
range of the species east of the Andes. A very distinctive new subspecies,
isolated in the upper Magdalena Valley, is described below.

Sporophila angolensis angolensis (Linnaeus)

Loxia angolensis Linnaeus 1766, Syst. Nat. 12th ed. 1:303. ‘‘Angola’” =
eastern Brazil.

Loxia torrida Scopoli 1769 Ann. I Hist. Nat. page 140. [Venezuela].

2Oryzoborus polinskii Sztolcman 1926, Ann. Zool. Mus. Pol. Hist. Nat.
5:230. Yurimaguas, Peru.

Characters.—Adult males with brown portions of underparts rich, deep
chestnut, rather than chocolate-brown.

Range.—Almost all of South America east of the Andes and south to
northern Argentina.

Specimens examined.—The majority of those in the institutions listed in
the acknowledgments.

Remarks.—Throughout most of the literature of the species, there have
been two races recognized east of the Andes: angolensis, which is stated
to range from central and eastern Brazil through eastern Bolivia, Paraguay,
and northern Argentina; and torrida, which is said to occur in Amazonian
Brazil, northeastern Peru, Colombia, Venezuela and the Guianas. Accord-
ing to Hellmayr (1938:246), torrida is ‘‘very similar to angolensis but slight-
ly smaller, the tail especially shorter, and with decidedly smaller, less bulky
bill.’ With the very large series available to me, I was unable to discern
any color differences in adult males and I could see no consistent difference
in the size of the bill. Furthermore, I found no significant difference in the
tail lengths of samples of adult males from the ranges of the two putative
subspecies (Table 1). Although the northern birds are very slightly smaller
on the average, there is broad overlap and it would be impossible to make
any distinction based on size. The single bird available from Paraguay had
a tail 58 mm long, which exceeds that of any of the others measured; it is
possible that the birds from the southernmost part of the range in Paraguay
and Argentina are larger, but there is insufficient material to ascertain this.

50) > PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Tail length (mm) of adult males of Sporophila angolensis.
n Range Mean
Eastern Brazil (angolensis) 6 52=56 54
Bolivia (angolensis) 5 51.5—56 53
Venezuela and Trinidad (“‘torrida’’) 6 51.5—53 oy
French Guiana (“‘torrida’’) 9 50.5—53.5 52

I cannot accept the validity of torrida and consider it to be a synonym of
angolensis.

There was some variation in the coloration of females and subadult males,
with certain individuals from Peru being considerably darker than typical
birds. There is a possibility that the name polinskii (Sztolcman) is valid.
However, I could find no logical pattern to the distribution of dark individ-
uals in Peru. Perhaps the molt patterns are different in portions of the Pe-
ruvian range of the species, and with better material it could be possible to
differentiate subspecies with specimens known to be in truly comparable
plumage. In any case, for the present I do not recognize any subdivision of
S. angolensis east of the Andes.

Sporophila angolensis theobromae, new subspecies

Holotype.—USNM 582224, adult male, collected 1 March 1968 at Melgar,
Tolima, Colombia, by C. J. Marinkelle (received in exchange from WFVZ).

Characters.—Adult males nearest S. a. angolensis but lower breast and
belly decidedly chocolate-brown, rather grayish, not chestnut.

Range.—Known only from two specimens from the upper Magdalena
River Valley in Tolima and Huila, Colombia.

Specimens examined.—Holotype, and a single additional adult male
(USNM 447682) from La Plata, Huila, Colombia, collected 14 April 1952 by
M. A. Carriker, Jr.

Etymology.—Of Theobroma, the genus of the cacao or chocolate tree
(literally ‘‘food of the gods’’ from Greek theos, god, and broma, food), in
reference to the coloration of the underparts in this bird.

Remarks.—This very isolated population is the only representative of the
angolensis group from west of the eastern Andes. Farther toward the mouth
of the Magdalena, only black-bellied individuals or intergrades are known
(see Olson, in press).

Acknowledgments

I am grateful to the following curators and institutions for lending, or
permitting examination of, the specimens cited above: Wesley E. Lanyon,

VOLUME 94, NUMBER 1 51

American Museum of Natural History, New York (AMNH); Frank B. Gill,
Academy of Natural Sciences of Phildelphia (ANSP); Kenneth C. Parkes,
Carnegie Museum of Natural History, Pittsburgh (CM); Melvin A. Traylor
and John W. Fitzpatrick, Field Museum of Natural History, Chicago (FM);
John P. O'Neill, Louisiana State University, Baton Rouge (LSU); Raymond
A. Paynter, Jr., Museum of Comparative Zoology, Cambridge, Massachu-
setts (MCZ); Lloyd F. Kiff, Western Foundation of Vertebrate Zoology,
Los Angeles (WF VZ). In addition, I employed the specimens in the National
Museum of Natural History, Smithsonian Institution (USNM). I am grateful
to Kenneth C. Parkes and Allan R. Phillips for their comments on the manu-
script. This is contribution Number 5 of the Wetmore Papers, a project
supported in part by trust funds from the Smithsonian Institution for com-
pleting unfinished work and study of undescribed material left by the late
Alexander Wetmore.

Literature Cited

Bowman, R.I. 1961. Morphological differentiation and adaptation in the Galapagos Finches.—
University of California Publications in Zoology 58:vii + 302 pages.

Hellmayr, C. E. 1911. A contribution to the ornithology of western Colombia.—Proceedings

of the Zoological Society of London 1911:1084—1213.

. 1938. Catalogue of Birds of the Americas. Part 11.—Field Museum of Natural History

Zoological Series 13(11):1-662.

Meyer de Schauensee, R. i966. The Species of Birds of South America and Their Distribu-

tion.—Livingston Publishing Company, Narberth, Pennsylvania. 577 pages.

. 1970. A review of the South American finch Oryzoborus crassirostris.—Notulae Na-

turae 418:1-6.

Olson, S. L. [In press.] Interaction between the two subspecies groups of the seed-finch
Sporophila angolensis in the Magdalena Valley, Colombia. Auk.

Paynter, R. A., Jr. 1970. Family Emberizidae. Jn R. A. Paynter, Jr., ed. Check-list of Birds
of the World. Volume 13.—Harvard University Press, Cambridge, Massachusetts. 443
pages.

Phelps, W. H., and W. H. Phelps, Jr. 1950. Seven new subspecies of Venezuelan birds.
Proceedings of the Biological Society of Washington 63:115—126.

Ridgway, R. 1901. The Birds of North and Middle America. Part I.—Bulletin of the United
States National Museum 50:1-715.

Sharpe, R. B. 1888. Catalogue of the Birds in the British Museum. Volume 12.—British
Museum, London. 871 pages.

Sick, H. 1963. Hybridization in certain Brazilian Fringillidae (Sporophila and Oryzoborus).—
Proceedings of the XIIIth International Ornithological Congress: 161-170.

Wetmore, A. 1957. The birds of Isla Coiba, Panama.—Smithsonian Miscellaneous Collections
134(9): 1-105.

National Museum of Natural History, Smithsonian Institution, Washing-
ton, D.C. 20560.

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 52-66

COMMENTS ON THE OSTEOLOGY OF BALISTOID
FISHES (TETRAODONTIFORMES), WITH NOTES
ON THE TRIODONTID PELVIS

James C. Tyler and Keiichi Matsuura

Abstract.—Two workers recently have independently described the os-
teological structure of cleared and stained specimens of a large number of
genera and species (ca. half the same) of fishes of the families Balistidae and
Monacanthidae (the superfamily Balistoidea, Tetraodontiformes). The phy-
logenetic and classificatory conclusions reached by Matsuura (1979) and
Tyler (1980) are rather similar, as are the majority of their osteological
observations. The few differences (ca. 5%) in their anatomical reports are
discussed herein.

Introduction

Matsuura (1979) published a detailed osteologically based phylogenetic
study of the balistoid fishes (Balistidae, triggerfishes; Monacanthidae, file-
fishes) utilizing cleared and alizarin stained specimens of 15 species repre-
senting 11 genera of balistids and 28 species representing 22 genera of mon-
acanthids. In his similarly based study Tyler (1980) used 21 species
representing 11 genera of balistids and 32 species representing 17 genera of
monacanthids. Matsuura’s coverage includes 57% of the balistids and 44%
of the monacanthids treated by Tyler, while Tyler’s coverage includes 80%
of the balistids and 50% of the monacanthids treated by Matsuura. Thus, of
the total of 70 species and 36 genera of balistoids treated by either Matsuura
or Tyler, 26 species and 25 genera are shared between the two (Table 1).
The species studied by Matsuura are mainly from Japan and the western
Pacific, while those used by Tyler are of more worldwide localities.

Since Matsuura and Tyler worked independently of one another, and are
of somewhat different (although cross-fertilized) cultural schools of ichthy-
ology, yet studied osteologically many of the same species and genera, an
opportunity to test the degree of similarity of their materials and observa-
tions presents itself. If one assumes that both researchers were careful in
their observations and precise in their descriptions, then similar results can
be presumed to be accurate in the vast majority of cases, for it is unlikely
that two attentive workers would frequently commit the same observational
error independently. Therefore, only the few differences in the osteological
descriptions of Matsuura and Tyler need to be discussed to clarify or rectify
these observations for the benefit of subsequent workers. The latter will

VOLUME 94, NUMBER 1 53

Table 1.—List of species studied (X) by Matsuura (1979) and Tyler (1980), and indication of
synonyms.

Examined by:

Species Matsuura Tyler

Balistidae

Abalistes stellatus
Balistapus undulatus x
Balistes capriscus —
Balistes forcipatus —
Balistes polylepis
Balistes vetula
Balistoides conspicillum
Balistoides viridescens
Canthidermis maculatus
Sufflamen bursa

= Hemibalistes bursa
Sufflamen chrysopterus

= Hemibalistes chrysopterus
Melichthys niger
Melichthys vidua
Odonus niger
Pseudobalistes flavomarginatus
Pseudobalistes fuscus
Rhinecanthus aculeatus
Rhinecanthus echarpe

= Rhinecanthus rectangulus
Rhinecanthus verrucosus
Sufflamen fraenatus
Sufflamen verres
Xanthichthys lineopunctatus
Xanthichthys mento
Xanthichthys ringens

See | p<escla|
[ee ee oi is Fe SASS Diabet dS Sl Det bia

pda i i |

xx «Kx

| x |
|

Monacanthidae

Acanthaluteres spilomelanurus
Acreichthys hajam
Alutera heudelotii
Alutera monoceros
Alutera schoepfi
Alutera scripta
Amanses scopas
Anacanthus barbatus

= Psilocephalus barbatus
Arotrolepis filicaudus
Brachaluteres trossulus
Brachaluteres ulvarum
Cantherhines dumerili
Cantherhines pardalis
Cantherhines pullus

pesca od? |
Dee pdinsd She ihc ii alec

Sea Ss
|

Kx x

54 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Examined by:

Species Matsuura Tyler

|
x

Cantherhines sandwichiensis
Chaetoderma penicilligera x x
= Chaetoderma spinosissimus
Eubalichthys mosaicus
Laputa cingalensis
Meuschenia freycineti
Meuschenia hippocrepis
Meuschenia trachylepis
Monacanthus chinensis
Monacanthus ciliatus
Monacanthus mylii
Monacanthus tuckeri
Navodon modestus
Navodon setosus
Navodon tesselatus
Nelusetta ayraudi
Oxymonacanthus longirostris
Paraluteres prionurus
Paramonacanthus barnardi
Paramonacanthus cryptodon
Paramonacanthus curtorhynchus
Paramonacanthus japonicus
Pervagor melanocephalus
Pervagor spilosomus
Pseudalutarius nasicornis
= Pseudaluteres nasicornis
Pseudomonacanthus peroni
Rudarius ercodes
Rudarius minutus
Scobinichthys granulatus
Stephanolepis auratus
Stephanolepis cirrhifer
Stephanolepis hispidus —-
Stephanolepis setifer —

ai SS ee S lx
||

<x «x >< |

Sea letbee || Sete at Salts Sees) ele Te Seb Sar See Le S|
esc <a <a eae |

peed Si Se lle a

have the opportunity to test in the arena of the 95% agreement between
Matsuura and Tyler what is here considered the likely assumption that they
are relatively careful reporters. It is here postulated that only about 5% of
the osteological observations of Matsuura and Tyler are in any kind of
conflict.

Even though Matsuura attempted a cladistically based analysis which
should exclusively use shared advanced character conditions (cladistic ap-
proach) to link groups in phylogenetic interpretations and generic segrega-

VOLUME 94, NUMBER 1 55

tions, while Tyler utilized the weighting and balancing of both generalized
and specialized features (gradistic or phenetic approach) to the same end,
the phylogenetic conclusions and generic recognitions between these two
osteologically based works are rather similar, with only a few notable ex-
ceptions (e.g., see Canthidermis, below).

The differences in the interpretative methodologies are not discussed fur-
ther here, with the following comments confined to the differential osteo-
logical observations. Most of these are accountable to statements about
genera based on different sets of species examined, or to different sizes of
individuals of the same species examined for what prove to be ontogenetic
features, and to individual variation between specimens of the same species.

Unless otherwise stated, references are to Matsuura (1979) and Tyler
(1980).

Balistidae

Frontal of Abalistes stellatus.—Matsuura states (p. 112, Fig. 40) that this
species is unique among the balistids in “‘having the frontal expanded greatly
. .. posteriorly beyond the level of the rounded postero-dorsal surface of
the cranium,’ while Tyler makes no such distinction. The explanation is
undoubtedly that Matsuura’s three specimens were all large adults (200.0-
230.2 mm SL) while Tyler’s single specimen was much smaller (87.6 mm
SL) and did not yet possess this posterolateral expansion of the frontal
which apparently develops only in larger adults. This region of the skull in
the 87.6 mm specimen is very similar to that of most other balistids.

Encasing scales of Balistapus undulatus.—Tyler states (1962, 1980) that
this species has but a single pair of scales in Segment III of the encasing
scale series at the posterior end of the pelvis, in contrast to two pairs in this
segment in Balistes [Verrunculus] polylepis, Balistes vetula, and other ba-
listids; Matsuura states (p. 73) that there are two pairs of scales there in B.
undulatus as well as in all other balistids. Matsuura examined 11 specimens
of 41.8 to 226.0 mm SL and Tyler examined 5 specimens of ca. 120 to 124.3
mm SL. Re-examination of Tyler’s material and of 3 additional specimens
(USNM 301981, 30.2 mm SL; USNM 197525, 46.0-68.7 mm SL) recently
cleared and stained shows Matsuura to be correct. However, there tends to
be fusion of the two pairs of scales in Segment III to the extent that it often
appears in large specimens that there is a single pair. Thus, in the recently
prepared 68.7 mm SL specimen, only a single pair of scales was evident
until all of the surface granulations and spiny processes had been shaved
away with a scalpel and the scale plates substantially thinned. Only then in
transmitted light was it possible to detect the closely apposed surfaces of
the two pairs, which even in this relatively small specimen were so consol-
idated that they nearly fused in places. Re-examination of the 124.3 mm SL
specimen previously reported by Tyler as having a single pair of scales in

56 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Segment III shows that this would be more accurately characterized as 11%
pairs (3 pieces), for the two pairs seem fully fused on one side but distinct
on the other. In the previously examined 121.6 mm SL specimen, both pairs
seem fully fused. It is obvious that Matsuura is correct in pointing out that
Segment III has two pairs of scales, at least developmentally, in all balistids,
even though some fusion between elements of the pairs can be expected in
some (usually larger) specimens of some species (such as B. undulatus).
Balistid postcleithrum.—Matsuura states (p. 78) that all balistids he ex-
amined have the postcleithrum as two pieces (dorsal and ventral postcleith-
ra), while (p. 80) all monacanthids have the postcleithrum as a single piece.
Tyler states (p. 101) that balistids have it as either a single piece or divided
into dorsal and ventral segments, in contrast to monacanthids in which the
postcleithrum is always a single piece. Tyler gives lateral view illustrations
(Figs. 70-72) of the entire skeleton of the balistids Rhinecanthus echarpe,
Sufflamen fraenatus, and Canthidermis maculatus showing the postcleith-
rum as a Single piece, in contrast to illustrations (Figs. 51, 69, 73) of Ba-
listapus undulatus, Balistes polylepis, and Xanthichthys lineopunctatus
showing the two segments of the postcleithra. Re-examination of the spec-
imens used by both Matsuura and Tyler indicates that Tyler is correct in
showing fusion of the postcleithrum into a single piece in some balistids.
Supposed scale bone of Canthidermis maculatus.—Matsuura describes
and illustrates (p. 109, Fig. 38) a scale bone as being present in this species
alone among all the balistoids, while Tyler (p. 43) says that a scale bone is
absent in all balistoids and in all other plectognaths (tetraodontiforms). This
is but one example among many of a bone that is present in such possibly
ancestral or sister group perciforms as acanthuroids and chaetodontoids but
lost by plectognaths in their largely reductive evolutionary processes. Mat-
suura’s two specimens were large adults (218.4 and 251.5 mm SL), while
Tyler’s was an especially nicely cleared 80.1 mm SL specimen, for which
a lateral view illustration of the entire skeleton is presented, and one larger
(216.9 mm SL) specimen. Tyler’s illustration (Fig. 72) shows a bone similar
to that thought to be a scale bone by Matsuura. Notes on the original draw-
ing state that this is simply the posterior portion of the sphenotic, which
superficially, on the lateral surface of the skull, is separated from the more
anterior portion of the sphenotic by a ventrad extension of the frontal. The
frontal makes slight contact with a dorsad extension of the pterotic and
apparently divides the sphenotic into an anterior and posterior portion which
are still fully continuous just below the surface. The 80.1 mm specimen has
been especially carefully re-examined to verify this point, and the ventrad
extension of the frontal that overlies the middle of the lateral surface of the
sphenotic laid back and excised on one side. This reveals (Fig. 1) the depres-
sion on the sphenotic that accommodates the extension of the frontal. The

‘n
~—

VOLUME 94, NUMBER |

supraoccipital epiotic

frontal

exoccipital

pterosphenoid

sphenotic prootic

‘ * . posttemporal

pterotic

basioccipital

parasphenoid Ist vertebra

Fig. 1. Posterior portion of cranium of Canthidermis maculatus, ANSP 100085, 80.1 mm
SL. Larger drawing above shows the downward flange of the frontal meeting the upward
extension of the pterotic, superficially dividing the sphenotic into forward and rear sections
which are clearly continuous below the surface as seen in the lower drawing in which the
downward flange of the frontal has been cut away (dashed line).

more posterior portion of the apparently divided sphenotic could be mis-
interpreted as a scale bone. Re-examination of the specimens used by Mat-
suura also shows the element described as a scale bone to be the rear of the
sphenotic.

This separation of the sphenotic by a superficial meeting of a ventrad
flange of the frontal and a dorsad flange of the pterotic also occurs to some
degree in at least large individuals of Balistes capriscus (Tyler, Fig. 74) with
partial external separation, in Sufflamen fraenatus (Tyler, Fig. 70) with
partial external separation, and in Balistes polylepis (Tyler, Fig. 69) with
external separation as complete as in Canthidermis.

Matsuura placed great emphasis on the supposed presence of a scale bone

58 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

in C. maculatus as an indicator of its primitiveness, and separated it off
first (Fig. 79) as the sister group of all other balistids on this basis. Fraser-
Brunner (1935:660—661) also thought Canthidermis to be the most general-
ized balistid, but only on the basis of it lacking enlarged tympanic scales
above the pectoral fin. Conversely, Tyler (p. 123) considered Canthidermis
to be only slightly less specialized than the most specialized of the balistids
(the Xanthichthys-Odonus line) and of about the same level of specialization
in its own way as Melichthys. Tyler considered Canthidermis specialized
for a more oceanic and pelagic existence than the other balistids (even more
so than Abalistes, an offshore form) with the scales of the body ‘‘all being
reduced in size and nonoverlapping [explaining the absence of distinctive
tympanic scales] and thus more flexible . . .”’ and ‘‘also specialized in hav-
ing the full ossification of the skeleton much delayed, as well as in having
the most rudimentary pelvic apparatus among the balistids,’’ including its
especially rudimentary pelvic fin ray element and relatively inflexible series
of encasing scales.

Balistoid urohyal shape.—Matsuura (p. 92 et seqg., Figs. 26, 28) describes
a difference between the urohyal in balistids (plate-like with several projec-
tions) and monacanthids (flattened and boomerang-like) overlooked by Ty-
ler in his comparative diagnoses of the two families.

Balistid epipleurals.—Matsuura states (p. 149) that the epipleurals in ba-
listids never extend posteriorly beyond the first caudal vertebra, while Tyler
(p. 167, Fig. 71) correctly shows that in Rhinecanthus they are attached as
far posteriorly as the fifth and sixth caudal vertebrae.

Balistoid posttemporal.—Both Matsuura and Tyler call attention to a fa-
milial distinction in the posttemporal between balistids and monacanthids,
but describe it with different emphasis. Matsuura states (p. 148-149) that
the posttemporal in balistids articulates with the epiotic, but that it does not
contact that bone in monacanthids. Tyler (p. 101, 135) states that the post-
temporal in balistids is held in a deep groove on the lateral surface of the
pterotic, but that in monacanthids it is held more superficially in a much
less deep groove on the lateral surface of the pterotic. Tyler makes no direct
statement on the relationship of the posttemporal to the epiotic. Matsuura
is correct in pointing out that the less sturdy posttemporal of monacanthids
is usually distinctly farther removed from the epiotic (least so in Oxymon-
acanthus, see Matsuura Fig. 47 and Tyler Fig. 122) than in balistids. In
some specimens of some species of balistids, however, it appears that the
posttemporal either does not make contact with the epiotic or barely does
so (see Tyler, Fig. 52, for Balistapus undulatus). It is excluded, or mostly
so, from that contact by the juncture of the sphenotic and pterotic behind
it, which intervenes between the posttemporal and epiotic. Perhaps there
are ontogenetic changes in the degree of exclusion. The proximity of the

VOLUME 94, NUMBER 1 59

posttemporal to the epiotic in balistids is sometimes closer as seen from the
inside of the cranium (cephalic vault) since the sphenotic and pterotic over-
lap the posterodorsal end of the posttemporal less extensively internally
than externally. In any case, the relationship of the posttemporal to the
epiotic is more of a quantitative than a qualitative distinction between the
two families.

Monacanthidae

Monacanthid scapular foramen.—Matsuura States (p. 80) that ‘‘the inner
surface of the scapula foramen is completely enclosed by the scapula in all
the monacanthid members, though the anterior edge of the outer surface of
the scapula foramen is surrounded by the cleithrum in these members: Ste-
phanolepis, Rudarius, Alutera, Oxymonacanthus, Pseudalutarius, and An-
acanthus,’’ whereas Tyler states (p. 172) that the scapular foramen is com-
pletely enclosed by the scapula in all monacanthids (as in balistids) except
Anacanthus and Pseudalutarius, two of the most specialized genera of the
family. Re-examination of the specimens used by Tyler, including the care-
ful disarticulation of the delicate scapula from previously undisturbed pec-
toral girdles, reaffirms that the scapula is incomplete around the foramen
(Fig. 2) anteriorly in A. barbatus and P. nasicornis, where the border of
the foramen is formed by the cleithrum. However, there is individual vari-
ation in the closure of the foramen, for re-examination of Matsuura’s spec-
imens (4 of P. nasicornis, 124.3—-150.2 mm SL; 2 of A. barbatus, 157.1-
206.4 mm SL), all of which are slightly to substantially larger than those
used by Tyler (2 of P. nasicornis, 50.8-108.3 mm SL; 3 of A. barbatus,
142.3-138.7 mm SL) shows the foramen sometimes to be complete.

Monacanthid pharyngobranchial and basibranchial elements.—Matsuura
extensively surveyed and amply illustrated the branchial arches in balis-
toids, probably more fully than ever before with any familial grouping of
fishes. Tyler’s interpretations of the branchial arches differ in only two
respects. Matsuura states (p. 102-103) that Oxymonacanthus longirostris
and Paraluteres prionurus are unique among the monacanthids in having a
toothless suspensory first pharyngobranchial in addition to the two toothed
elements found in all other monacanthids, and that P. prionurus is also
unique in having four basibranchials rather than the three of all other mon-
acanthids (except Anacanthus with only two). By omission to the contrary,
Tyler implies that both of these species have the normal monacanthid bran-
chial arrangement of only two pharyngobranchials (both toothed) and three
basibranchials.

Re-examination of Tyler’s specimens of these two species (8 of O. lon-
girostris and | of P. prionurus) indicates that none of them has a first tooth-
less suspensory pharyngobranchial and that P. prionurus has only three

60 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

cieithrum/

scapula

scapula

~ “\coracoid

Fig. 2. Medial view of left scapula of (A) Pseudalutarius nasicornis, USNM 294075, 50.1
mm SL, and (B) Anacanthus barbatus, ANSP 109648, 142.3 mm SL, in contrast to the more
normal condition of the scapula (C) as seen, for example, in Alutera monoceros, HUMZ 38062,
155.2 mm SL, in lateral view.

basibranchials, as do most other monacanthids. Re-examination of Mat-
suura’s specimens (2 of O. longirostris and 2 of P. prionurus) shows that
the element described as the toothless suspensory first pharyngobranchial
is actually a distal part of the first epibranchial. This distal part is abnormally
separated from the proximal part of the first epibranchial by an unossified
portion. Examination of additional specimens of O. longirostris (HUMZ
40658, 50366) confirms the normal pharyngobranchial arrangement in this
species, with two pharyngobranchial elements only, both toothed. Re-ex-
amination of Matsuura’s specimens of P. prionurus indicates that the bone
interpreted as the fourth basibranchial is actually the anterior portion of the
fourth ceratobranchial, abnormally fused into a shaft-like element and sep-
arated from its posterior region by an unossified segment.

To avoid confusion, it should be noted that there is a typographical error
in Table 4 (p. 108) of Matsuura’s work, for Paraluteres is listed as having
the normal two pharyngobranchials, while it is described in the text (p. 108)
and illustrated (Fig. 34) as having three pharyngobranchials.

Posttemporal of Anacanthus barbatus.—Tyler states (p. 167) that the
‘‘posttemporal is present in all monacanthids except Psilocephalus barba-
tus, none of the three adult specimens of which shows any evidence of one.
Whether the ossification center for the posttemporal is lost or incorporated
indistinguishably with that of the pterotic remains problematical,’’ while
Matsuura (p. 116) has been able to distinguish a bump on the pterotic in his
specimens [Anacanthus| which may represent the fused posttemporal.

Dewlap expansion in monacanthids.—Matsuura states (p. 77) that “*In

VOLUME 94, NUMBER I 61

contrast with the primitive monacanthids, the advanced monacanthids such
as the fishes of the genus Brachaluteres can only slightly extend downward
the reduced pelvis without the incasing scales, however, they have the other
mechanism to inflate their abdomen’”’ [=inflatable stomach]. This statement
was made to explain the functional significance of the stomach of Brachal-
uteres. However, it should be borne in mind that one of the most highly
specialized of all monacanthids, the exceptionally elongate Anacanthus,
which is like the deep bodied Brachaluteres in its very weakly developed
pelvis without encasing scales, can flare one of the largest dewlaps among
all of the monacanthids, generalized and specialized. The ability to flare a
dewlap is of little phylogenetic significance.

Third tooth of dentary in monacanthids.—Matsuura states (p. 83) that the
third or posteriormost (last) tooth in the dentary “‘is extremely reduced in
size” in Brachaluteres (ulvarum examined) and Pseudomonacanthus (per-
oni), and absent altogether in Rudarius (ercodes), Oxymonacanthus (lon-
girostris), Paraluteres (prionurus), and Anacanthus (barbatus). Tyler
states (p. 173) that the third tooth of the dentary is much smaller than the
others in Pseudalutarius (nasicornis), and absent altogether in Rudarius
(ercodes and minutus), Oxymonacanthus (longirostris examined, and pre-
sumably in halli also), Paraluteres (prionurus), Anacanthus (barbatus),
and Brachaluteres (trossulus examined, and presumably in baueri and wol-
fei also), while Tyler did not examine Pseudomonacanthus. It seems ob-
vious on the basis of these two reports that the development of the third
dentary tooth is variable within Brachaluteres, being reduced in size in the
species (ulvarum) studied by Matsuura and absent in that (trossulus) studied
by Tyler or reported in the literature (baueri and wolfei), and that it is
absent in all species of Rudarius, Oxymonacanthus, Paraluteres and Prion-
urUus.

This leaves in conflict the reported size of the third dentary tooth of the
monotypic Pseudalutarius (nasicornis). Matsuura shows it (Fig. 22) to be
of relatively large size in the illustrated individual among his four specimens
of 124.3-150.2 mm SL, whereas Tyler reports this tooth as of much reduced
size in his two smaller specimens (50.8—108.3 mm SL). As a check on vari-
ability in the size of the third dentary tooth in Pseudalutarius nasicornis,
an additional specimen has been cleared and stained (USNM 294075, 50.1
mm SL), and it too has the third dentary tooth much reduced in size (Fig.
3, left) as reported by Tyler. On the other hand, re-examination of Mat-
suura’s specimen of this species (HUMZ 38074, 150.2 mm SL) confirms that
the third tooth is relatively larger (Fig. 3, right). There obviously is variation
in the size of the third dentary tooth in this species, and it may increase
with increasing specimen size.

Ribs of Pseudalutarius nasicornis.—Matsuura describes (p. 140) the rib-

62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Lateral view of teeth in left dentary of Pseudalutarius nasicornis, USNM 294075,
50.1 mm SL (left) and HUMZ 38074, 150.2 mm SL (right), to show the variation in size of the
third (posteriormost) tooth; anterior to left.

like element attached to the first vertebra as an epipleural, and so illustrates
it (Fig. 56). Conversely, Tyler (p. 167) states that the element attached to
the first vertebra is a pleural rib like those that follow it on the other ab-
dominal vertebrae, and so illustrates it (Figs. 114, 123). Re-examination of
Matsuura’s four specimens and of an additional recently cleared and stained
specimen (USNM 294075, 50.1 mm SL) indicates that the element attached
to the first vertebra should be recognized as a pleural rib rather than an
epipleural, for the following reasons: 1) the element is longer than the epi-
pleurals that are attached to the pleural ribs of the second to seventh ver-
tebrae; 2) the element and the pleural ribs are distinctly more ventrally
directed in position than are the epipleurals; 3) the element is in positional
Orientation and symmetrical series with the other pleural ribs (second to
seventh vertebrae); 4) the element courses along the peritoneal wall of the
abdominal cavity like the other pleural ribs, rather than, as with the series
of epipleurals, being in the septum between the epaxial and hypaxial mus-
culature.

Matsuura also states (Table 9, p. 139) that the epipleurals in monacanthids
other than Pseudalutarius (including in Rudarius ercodes and Brachalu-
teres ulvarum), start on the second abdominal vertebra, while Tyler (p. 167)
says and illustrates that they usually start from the second abdominal ver-
tebra, but that they start from the third vertebra in Rudarius ercodes and
R. minutus as well as in the single species he examined of Brachaluteres
(trossulus). Re-examination of Matsuura’s specimens of R. ercodes and B.
ulvarum confirms Matsuura’s statements. Thus, there is occasional variation
between species and between individuals of the same species in the place

VOLUME 94, NUMBER I 63

of origin of the first epipleural, variously from the second abdominal ver-
tebra (usually) or from the third (more rarely).

Second dorsal spine of Brachaluteres.—Matsuura states (p. 126) that all
monacanthids have two dorsal spines, except Anacanthus which has only
one, while Tyler says (p. 146) that both Brachaluteres and Anacanthus have
only a single dorsal spine. The conflict is between the species studied of
Brachaluteres, Matsuura examining B. ulvarum, in which a second spine
is present, and Tyler examining B. trossulus, in which it is absent. Tyler
should have known this, for he had radiographed the two co-types of B.
ulvarum (Stanford University 7128, now California Academy of Sciences),
as reported in his table of plectognath vertebral counts, and the radiographs
clearly show a small but normally developed second spine to be present.
The second spine is absent in two cleared and stained and 20 radiographed
specimens of B. trossulus studied by Tyler.

Upper free hypural in monacanthids.—Matsuura states (p. 147) that an
upper free hypural is absent in Rudarius ercodes, Brachaluteres ulvarum,
and in adult Alutera scripta. Tyler reports (p. 173, 179, et seq.) the upper-
most hypural to be free in Alutera scripta and the other three species of
Alutera he examined (heudelotii, monoceros and schoepfi) as well as in
Brachaluteres trossulus, but to be absent in most specimens studied of
Monacanthus ciliatus, both species studied of Rudarius (ercodes and mi-
nutus), and in Amanses scopas. Matsuura describes (p. 147) and illustrates
(Fig. 66) an upper free hypural in a juvenile Alutera scripta and its absence
(through fusion) in an adult. Matsuura’s four specimens of A. scripta were
59.0—295.8 mm SL and the two of Tyler were 46.2—73.3 mm SL. Fusion of
the upper free hypural obviously occurs with increasing specimen size in
A. scripta and in other species of monacanthids.

Palatine of Anacanthus barbatus.—Matsuura emphasizes (p. 88) the
unique close articulation of the palatine with the ectopterygoid in Anacan-
thus barbatus, which is a more distant articulation mediated by a strong
ligament of varying length in all other monacanthids. Both Matsuura (Fig.
25) and Tyler (Figs. 110, 120) illustrate this close apposition of articulation
in A. barbatus, but Tyler does not comment on it in the text.

Pelvis-pectoral girdle articulation in Pervagor.—Tyler (p. 180, Fig. 105)
comments on a unique specialized feature of the pelvis in Pervagor not
mentioned by Matsuura, this being the development on the anterior region
of the pelvis, just behind the pectoral girdle, of a lateral knob-like expansion
which articulates with a similar expansion on the posterior edge of the cor-
acoid, forming a point of pivot around which the pelvis rotates.

Encasing scales of Chaetoderma penicilligera.—Matsuura states (p. 59)
that there are three series of encasing scales at the end of the pelvis in C.
penicilligera and so illustrates it (Figs. 7-8), while Tyler describes (p. 178,
Fig. 88) only two series of encasing scales in this species. Examination of

64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Segment Il

Segment Ill

Segment |

Fig. 4. Lateral view of encasing scales at end of pelvis (anterior to left) of Chaetoderma
penicilligera, HUMZ 40645, 172.3 mm SL.

an additional specimen of C. penicilligerum (HUMZ 40645, 172.3 mm SL)
clearly shows the encasing scales in three series (Fig. 4), although the sec-
ond segment is not broadly present on the ventral surface of the apparatus.

Triodontidae

Pelvis of Triodon.—Matsuura presents an extensive survey of the pelvic
apparatus of balistoids, greatly expanding on the coverage provided by Tyler
(1962, 1980), especially for the rudimentary fin ray element. The latter lies
buried beneath encasing scales and is difficult to dissect free for detailed
study and illustration. As an aside to that exposition on the balistoid pelvic
apparatus, Matsuura describes the pelvis of Triodon, the most generalized
Recent member of the gymnodont line of plectognaths. Matsuura states (p.
78) that there is a ‘“‘very small cartilage plug which is quite similar to the
structure found in the advanced monacanthids”’ although ‘‘it is very difficult
to locate since the plug is tightly enclosed by tough connective tissue at the
end of the posterior part of the pelvis.’’ In Tyler’s descriptions (1962, 1980)
of the pelvis in 7Jriodon no mention is made of these or of most other
cartilaginous elements of the skeleton.

Re-examination of one (ANSP 98917, 463.3 mm SL) of the two specimens
of Triodon previously studied by Tyler shows that cartilaginous plugs exist
at both the anterior and posterior ends of the pelvis (Fig. 5). The plugs
surround the paired ends of the pelvis, as one would expect in an endo-
chondral bone of a plectognath. Each plug is bilaterally paired, with the two
halves of the plug bound together by connective tissue and terminating both
anteriorly and posteriorly the ossifications of the pelvis (which two halves
are fused together in much of the posterior half of their lengths but not at

VOLUME 94, NUMBER 1 65

lateral

nee = dorsal peeaerenanna se
ee a dorsal
a pelvis

pelvis

cartilage

Fig. 5. Pelvis of Triodon macropterus, ANSP 98917, 463.3 mm SL. Above, lateral view of
the entire pelvis (178 mm length); middle, dorsal views of the anterior and posterior ends of
the pelvis; below, enlargements of the dorsal views of the anterior and posterior ends of the
pelvis.

the extreme posterior end). The structure of the paired plugs is basically
similar at both ends of the pelvis, although far larger at the anterior end.
Therefore, Tyler does not believe that the paired plug at the posterior end
of the pelvis in Triodon is necessarily homologous to the far more complex
and less distinctly paired plug supporting the highly modified fin ray element
in balistoids.

Examination of an additional specimen of the rarely collected Triodon
(NSMT-P 3415, 370.0 mm SL) by Matsuura shows the same structure of the
pelvis and cartilages as in Tyler’s specimen. However, Matsuura believes
that the plug at the posterior end of the pelvis of Triodon is possibly ho-
mologous to the cartilaginous plug of the fin ray element of balistoids for
the following reasons: 1) the condition of the plug in Triodon is closely
similar to that of such advanced monacanthids as Pseudalutarius and
Oxymonacanthus, except for being paired; 2) the paired nature of the plug
in Triodon 1s related to the divided condition of the pelvis, while the single
plug of monacanthids corresponds to the undivided pelvis of that family; 3)
the advanced monacanthids have a cartilaginous region at the anterior end
of the pelvis similar to that of Triodon.

Only detailed histological analysis of the ontogenetic development of
these cartilagenous plugs in Triodon and balistoids will solve the question
of their possible homology.

Summary

The majority of the osteological observations that were made indepen-
dently by Matsuura (1979) and Tyler (1980) on a largely similar coverage of

66 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

genera and species of balistoids are compatible. Most of the differences are
due to statements made about genera based on different sets of species
examined or about species based on different sized specimens examined,
and on other intraspecific variation.

Acknowledgments

We are grateful to Daniel M. Cohen and Bruce B. Collette of the National
Marine Fisheries Service, National Systematics Laboratory, for providing
space and equipment for microscopic examination of specimens, and to
William Aron, National Marine Fisheries Service, Office of Marine Mam-
mals and Endangered Species, and Frank B. Golley, National Science Foun-
dation, Division of Environmental Biology, for the research time of one of
us (JCT). Our thanks also go to Victor G. Springer, U.S. National Museum
(USNM) for permission to clear and stain a number of specimens mentioned
here, and to Takao Igarashi and Kunio Amaoka, Laboratory of Marine
Zoology, Faculty of Fisheries, Hokkaido University (HUMZ), William
Smith-Vaniz, Academy of Natural Sciences of Philadelphia (ANSP) and
William N. Eschmeyer, California Academy of Sciences (specimens from
Stanford University, SU) for the loan of materials. Ryoich Arai, Department
of Zoology, National Science Museum, Tokyo (NSMT-P), Joseph Russo,
National Systematics Laboratory, Richard Vari, U.S. National Museum,
and Richard Winterbottom, Royal Ontario Museum, offered valuable sug-
gestions to improve the manuscript.

Literature Cited

Fraser-Brunner, A. 1935. Notes on the plectognath fishes.—I. A synopsis of the genera of the
Family Balistidae.—Ann. Mag. Nat. Hist. (10)15:658-663.

Matsuura, Keiichi. 1979. Phylogeny of the Superfamily Balistoidea (Pisces: Tetraodonti-
formes).—Mem. Fac. Fish., Hokkaido Univ. 26(1/2):49-169.

Tyler, James C. 1962. The pelvis and pelvic fin of plectognath fishes; a study in reduction.—

Proc. Acad. Nat. Sci. Philadelphia 114(7):207—250.

. 1980. Osteology, phylogeny and higher classification of the fishes of the Order Plec-

tognathi (Tetraodontiformes).—U.S. Dep. Commerce, Nat. Ocean. Atmos. Admin.,

Nat. Mar. Fish. Serv., NOAA Tech. Rep., NMFS Circ. 434, 422 pp.

(JCT) Division of Environmental Biology, National Science Foundation,
Washington, D.C. 20550, U.S.A. (Research Associate, American Museum of
Natural History, Department of Ichthyology, and U.S. National Museum of
Natural History, Department of Vertebrate Zoology), and (KM) Depart-
ment of Zoology, National Science Museum (Natural History Institute),
3-23-1 Hyakunin-cho, Shinjuku-ku, Tokyo 160, Japan.

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 67-75

A NEW POISON-DART FROG (ANURA:
DENDROBATIDAE) FROM THE FOREST OF
SOUTHEASTERN COLOMBIA

William F. Pyburn

Abstract.—Dendrobates myersi, a new species of poison-dart from from
the Colombian rain forest, is related to members of the Dendrobates fe-
moralis species group. The new frog is terrestrial, diurnal, and probably
territorial. Its call is a long series of double-noted chirps given by isolated
individuals. In captivity D. myersi is aggressive toward other dendrobatids.
The larva and reproductive characteristics of the new species are unknown.

Early in August 1972, near the Cacua Village of Wacara, I listened to the
penetrating notes of a dendrobatid frog calling from the leaf litter of the
forest floor. The call was a long series of double-noted chirps, unlike that
of any frog I had heard before. I recorded the call with a Uher 4000 Report
L tape recorder, and a Cacua Indian companion captured the brightly pat-
terned frog where it sat at the base of a low shrub. The time was about one
hour before sunset.

Since that initial encounter, I have obtained six additional specimens of
this unusual frog, one of which was sent alive to Charles Myers of the
American Museum of Natural History. Analysis of the skin gland secretions,
to be reported by Myers elsewhere, together with the color pattern and
unusual, apparently territorial call, show that these seven specimens rep-
resent an unnamed species of poison-dart frog.

Dendrobates myersi, new species
Rigg |

Holotype.—UTA A-3989, adult male collected near Wacara (elev. 216 m,
long. 69°53’W, lat. 1°08’N), Comisaria de Vaupés, Colombia, 2 Aug. 1972
by J. K. Salser, Jr., Vincente Lopez, Arquelao Gallego, and W. F. Pyburn.

Paratypes.—UTA A-7634, juvenile collected at type-locality, 3 Aug. 1972
by J. K. Salser, Jr. and W. F. Pyburn; AMNH 104681, juvenile collected
at type locality, 3 Aug. 1972 by J. K. Salser, Jr. and W. F. Pyburn; AMNH
102173 adult male (skinned carcass) obtained 21 April 1976 by Pyburn at
Yapima (69°28’W, 1°03’N), Comisaria de Vaupés, Colombia; AMNH
104682, adult female obtained 5 Aug. 1974 by Nathan Waltz at Yapima;
AMNH 104683, adult male obtained 2 Sept. 1975 by Nathan Waltz near

68 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Dendrobates myersi, paratype, AMNH 102173. SVL = 29.5 mm. From a color
transparency by C. W. Myers.

Yapima; FM 1331, adult male collected 7 April 1979 by William W. Lamar
near Mitu, Comisaria de Vaupés, Colombia.

Diagnostic characters.—A moderate sized Dendrobates with a snout-
vent length in males of about 28 mm. Combinations of the following char-
acteristics distinguish Dendrobates myersi from all other dendrobatid frogs:
maxillary and premaxillary teeth present; toes basally webbed; first finger
longer than second; tarsal tubercle and ridge present; omosternum present;
no light dorsolateral stripe, sides black; a complete light ventrolateral stripe;
dorsum brown; bright yellow flash colors in groin and on lower arm surface;
voice a long series of double-noted chirps.

In general appearance, Dendrobates myersi resembles D. femoralis (Fig.
2) and D. zaparo of the femoralis species group of Silverstone (1976). Den-
drobates myersi differs from Dendrobates femoralis in having an indistinct,
or no, light dorsolateral stripe separating the brown back color from the
black of the sides. D. femoralis has a complete, well defined, dorsolateral

VOLUME 94, NUMBER 1 69

Fig. 2. Dendrobates femoralis from Pebas (Loreto) Peru, AMNH 103570. SVL = 26 mm.
From a color transparency by C. W. Myers.

stripe (Boulenger, 1883; Cochran and Goin, 1970) that is said by Silverstone
(1976) to be yellow or golden color in life. I have seen color transparencies
of live D. femoralis from Amazonian Peru (Loreto, Pebas) and from the
Coppername River of Surinam (Saramacca, Raleigh Cataracts). The Peru-
vian specimen showed a pale cream-colored dorsolateral stripe and the one
from Surinam showed a golden yellow stripe. There is also a difference
between the two species in the marking on the thigh. In life, D. femoralis
has, on the proximal part of its upper thigh surface, a sharply defined red
or yellow diagonal mark that is not present in D. myersi. The upper thigh
surface of D. myersi may be red, but the red area, when present, is distal
and its leading edge is diffuse so that it does not form a sharply defined
mark.

Dendrobates myersi differs from D. zaparo in the color and texture of the
dorsal skin. In life the dorsum of D. zaparo is red (Silvertone, 1976), where-
as live D. myersi has a brown dorsum, sometimes with evenly distributed
dark reticulations or spots. Although the dorsum of both species is granular,
the granules of D. zaparo are more densely packed together and more nu-
merous than the granules of D. myersi. In preserved specimens, the dorsal

70 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

granules of D. zaparo are pale on a dark ground color, whereas the reverse
is true of D. myersi.

With the possible exception of Dendrobates tricolor, all of the species in
the femoralis group are much smaller than Dendrobates myersi. In preser-
vative, D. tricolor, unlike myersi, has a yellow dorsum and a yellow or
white dorsolateral stripe. The small species may further be distinguished
from D. myersi as follows: D. anthonyi has light middorsal and dorsolateral
Stripes (absent in myersi); D. boulengeri has in life a complete yellow or
red dorsolateral stripe (Silverstone, 1976) that fades to cream or white in
preservative, and has an irregular, reticulated or spotted pattern on the
upper thigh surface (both absent in myersi); D. espinosai is in life a red-
backed frog (Funkhouser, 1956) that in preservative has a dark brown dor-
sum, no ventrolateral light stripe and sometimes an incomplete dorsolateral
light stripe (D. myersi has a medium to light brown dorsum and a white
ventrolateral stripe).

Description (five adults including the holotype).—A medium sized den-
drobatid (Table 1) with teeth on the premaxillary and maxillary bones. Vocal
slits are present in males, absent in females. The tongue is free laterally and
posteriorly, and has prominent papillae on the lateral and anterior parts of
its upper surface. Dorsal skin of the head, back and shank is granular;
ventral and lateral skin surfaces are smooth. Some specimens have a poorly
defined, glandular, dorsolateral ridge.

The lower eyelid is transparent. Males have two longitudinal pleat-like
folds in the skin of the gula. The posterodorsal rim of the tympanum is
concealed. There are basal webs between toes II and III and between III
and IV. The tarsal tubercle 1s connected by a dermal ridge to a prominent
inner metatarsal tubercle. The outer metatarsal tubercle is small but distinct
and there are large, ovoid subarticular tubercles on the toes. On the hand,
a large palmer tubercle is about twice the size of the thenar tubercle and
there are well developed, rounded subarticular tubercles on the fingers. The
first finger is longer than the second. An omosternum is present and the
muscle tissue is pink.

In life the skin of the dorsum is medium to light brown with closely
spaced, darker brown granules. The sides of the head and body are blue-
black and there is no light dorsolateral stripe separating the dorsal color
from the color of the side. The iris is golden, densely suffused with black.
There is a pale cream ventrolateral stripe passing from below the nostril,
along the upper lip and side of body, to the groin, where it becomes bright
yellow. The black of the side extends to the anterior base of the thigh and
around the posterior end of the body to, or nearly to, the vent. The blue-
grey to dark brown of the gula and chest merges with dark reticulations and/
or spots on the otherwise cream-white abdomen. Lower surfaces of the hind
limbs are generally marked like the skin of the abdomen. The axilla, pos-

71

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0.1 0.2 0.3 0.4 0.5 0.6 0.7 08 0.9 1.0 1.1 1.2 1.3 1.4
SECONDS

Fig. 3. Sound spectrogram (narrow band filter) of 6 notes (3 couplets) of Dendrobates
myersi from a recording of the holotype at the type locality, 2 August 1972, air temp. ~ 24°C.

terior surface of the upper arm, and anterior thigh surface are yellow. There
is an irregular black band or group of black spots on the posterior thigh
surface extending from the bend of the knee toward the vent; above this the
thigh is reddish brown to coral red. The upper surface of the shank is me-
dium brown with darker granules much like the dorsum. There is an irregular
dark brown spot or stripe in the bend of the tarsus, and the fingers and toes
may bear brown spots.

There is no sexual dimorphism in color pattern. The various color mark-
ings of the female are duplicated by one or more of the males.

Color of juveniles.—Two juveniles in the type series have the color pat-
tern of the adults, except that the limbs of the young frogs are straw-colored
to creamy pink and are unmarked.

Voice.—The call (Fig. 3) is a long series of whistle-like notes sometimes
continuing for a minute or more if the caller is not disturbed. The notes are
‘chirp calls’? (Myers and Daly, 1976), but unlike the chirps of other den-
drobatids they are uttered in couplets or note pairs.

Two frogs at Wacara, timed with a stop watch, called at a rate of 228.6
notes per min (160 notes, air temperature about 24°C). Sound spectrograms
show a dominant frequency band between 3100 and 3800 Hz and a note-pair
duration of about 0.5 s.

Dendrobates espinosai is the only member of the femoralis group whose
voice has been recorded and published as a sound spectrogram (Myers and
Daly, 1976: Fig. 23). The voice of D. espinosai is a “‘buzz’’ consisting of
a uniform series of pulses produced too rapidly to be resolved by the human
ear. The call of D. espinosai does not resemble the easily resolved double
note chirps of D. myersi (Fig. 3). Several dendrobatids produce calls con-
sisting of evenly spaced chirps (Myers and Daly, 1976) but D. femoralis is

VOLUME 94, NUMBER 1 3

the only other species known to produce a double noted call. Duellman
(1978) described the call of D. femoralis as **. . . a series of high-pitched,
quickly repeated pairs of notes: ‘peep-peep, peep-peep, peep-peep.’’’ The
apparent similarity in call and in morphology between D. femoralis and D.
myersi Suggests that these two species are closely related.

Natural history.—Dendrobates myersi is terrestrial and, like other den-
drobatids, it is diurnal. Males call from isolated positions in the leaf litter
on the forest floor. The frogs are not common and are always well separated
from each other (10 m or more) when calling, suggesting that the call is a
territorial advertisement. Myers (personal communication) observed a cap-
tive adult male D. myersi behaving aggressively toward dendrobatids of
other species (Phyllobates aurotaenia, P. vittatus and P. terribilis) in the
same terrarium. The D. myersi gave a short series of ‘“‘peeps’’ and hopped
on the other frogs. At other times it called from a certain, seemingly pre-
ferred, position in the terrarium.

In nature males of D. myersi answer to playback recordings of their calls.
Most of the specimens were active when collected, but one (AMNH 104683)
was found on a rainy day under cut palm leaves.

The larva and reproductive characteristics of myersi are unknown.

Remarks.—Dendrobates myersi is presently known only from the valley
of the Vaupés River and its tributaries in southeastern Colombia. The In-
dians of the region (Cacua, Cubeo, Guanano) are familiar with this species,
but they do not fear it and apparently do not use its skin secretions to poison
their blow gun darts.

Myers et al. (1978) reassigned several species to the genus Dendrobates
that were included in the genus Phyllobates by Savage (1968), Silverstone
(1976), Cochran and Goin (1970), and others. Among the species reassigned
to Dendrobates were members of Silverstone’s femoralis group, to which
the new species apparently belongs. I have tentatively accepted the arrange-
ment of Myers et al. (1978) and have therefore placed the Vaupés species
in the genus Dendrobates.

Etymology.—This species is named for Charles W. Myers of the Ameri-
can Museum of Natural History in recognition of his innovative research in
dendrobatid systematics.

Specimens Examined

Abbreviations: AMNH, American Museum of Natural History, New
York; FM, Federico Medem private collection, Villavicencio, Colombia;
USNM, JAP, National Museum of Natural History, Washington, D.C.;
UTA, University of Texas at Arlington, Arlington, Texas; CWM, Charles
W. Myers field series.

74 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Dendrobates anthonyi Noble—ECUADOR: El Oro, 3 km E Pasaje, 30 m
(AMNH 89630-635).

Dendrobates boulengeri Barbour—COLOMBIA: Narino, Isla Gorgona,
Gorgonilla (AMNH 62892-93), near prison camp (USNM 145248, 145249-
2k

Dendrobates espinosai Funkhouser—ECUADOR: Pichincha, Rio Baba,
5-10 km SSW Santo Domingo de las Colorados, 500 m (AMNH 89671-76),
Centro Cientifico Rio Palenque, 200 m (CWM 16021-—024); Rio Blanco, near
mouth of Rio Yambi, about 700 m (USNM 204506-10).

Dendrobates femoralis Boulenger—BRASIL: Amazonas, Igarape Puru-
zino, Rio Madeira (USNM 201861-63), Curuca, Rio Madeira (USNM
201864); GUYANA: Kartabo (AMNH 39679-80); PERU: Loreto, Centro
Union, lower Rio Aucayo, 20 km SE Iquitos, 3°48’S, 73°3’W (AMNH
88526—28).

Dendrobates tricolor Boulenger—ECUADOR: Azuay, Rio Minas, 20 km
W Santa Isabel on Cuenca-Machala Rd, 1250 m (USNM 166897, JAP 3600-—
01, 3606-08), ca. 16 km W Santa Isabel, 1000 m, Rio Jubones drainage
(CWM 16025-029).

Dendrobates zaparo Silverstone—ECUADOR: no specific locality
(AMNH 52881); Pastaza, 0.5 NE Puzo, 985 m (USNM 204534); Vera Cruz,
about 10 km E Puzo, 1016 m (USNM 204544-45); Cabeceras del Rio Bo-
bonaza, Camino a Guamajaco, 677 m (USNM 204546); 2 km S Shell Mera,
985 m (USNM 204547); PERU-ECUADOR frontier: Gauches (on Rio Pas-
taza) (AMNH 52882).

Acknowledgments

I am grateful to Charles W. Myers for permission to examine his color
transparencies and notes, and for the loan of specimens under his care. Also
for the loan of specimens I am indebted to W. Ronald Heyer and Ronald
Crombie (National Museum of Natural History), and William W. Lamar and
Federico Meden (Instituto Ciencias Naturales, Villavicencio, Colombia).
Officials of Inderena gave permission to collect specimens in Colombia. My
field companions at various times were Wanda Carl Pyburn, J. K. Salser,
Jr., Vicente Lopez, Arquelao Gallego, Lois Lores, Marilyn Cathcart, and
Nathan Waltz. Beverly Hood typed the manuscript. Travel to and from
Colombia, and in Colombia, was financed by the American Philosophical
Society (Johnson Fund) and by the University of Texas at Arlington (Or-
ganized Research Fund).

Literature Cited

Boulenger, G. A. 1883. Ona collection of frogs from Yurimaguas, Huallaga River, Northern
Peru.—Proc. Zool. Soc. London IV:635-638.

VOLUME 94, NUMBER 1 WS

Cochran, D., and C. J. Goin. 1970. Frogs of Colombia.—Bull. U.S. Nat. Mus. 288:1-655.

Duellman, W. E. 1978. The biology of an equatorial herpetofauna in Amazonian Ecudaor.—
Univ. Kansas Mus. Nat. Hist. Misc. Publ. 65:1-352.

Funkhouser, J. W. 1956. New frogs from Ecuador and Southwestern Colombia.—Zoologica
41:73-80.

Myers, C. W., and J. W. Daly. 1976. Preliminary evaluation of skin toxins and vocalizations
in taxonomic and evolutionary studies of poison-dart frogs (Dendrobatidae).—Bull.
Amer. Mus. Nat. Hist. 157:175—262.

——s , and B. Malkin. 1978. A dangerously toxic new frog (Phyllobates) used by
Embera Indians of western Colombia, with discussion of blowgun fabrication and dart
poisoning.—Bull. Amer. Mus. Nat. Hist. 161:309-365.

Savage, J. M. 1968. The dendrobatid frogs of central America.—Copeia 1968:745-776.

Silverstone, P. A. 1975. A revision of the poison-arrow frogs of the genus Dendrobates

Wagler.—Nat. Hist. Mus. Los Angeles County, Sci. Bull. 21:1—5S.

. 1976. A revision of the poison-arrow frogs of the genus Phyllobates Bibron in Sagra

(family Dendrobatidae).—Ibid. 27:1-53.

Department of Biology, The University of Texas at Arlington, Arlington,
Texas 76019.

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 76-87

ON DORVILLEIDAE AND IPHITIMIDAE
(ANNELIDA: POLYCHAETA) WITH A REDESCRIPTION
OF ETEONOPSIS GERYONICOLA AND A
NEW HOST RECORD

Gary R. Gaston and David A. Benner

Abstract.—The symbiotic polychaete, Eteonopsis geryonicola Esmark
(Dorvilleidae, Polychaeta), occurs in the branchial chambers of the crabs
Geryon tridens Krgyer, Geryon quinquedens Smith, and Cancer borealis
Stimpson in the North Atlantic. The latter is a new host record. The species
is herein redescribed and the original genus, Eteonopsis, revived. A histor-
ical summary is provided to clarify confusion concerning the original de-
scription. The family, Iphitimidae, a similar group of branchiate poly-
chaetes, are synonymized with Dorvilleidae. Eteonopsis occurs with
significantly greater frequency and intensity in male than female crabs. Gut
analyses of the worms failed to discern their mode of nutrition. A key to
the Dorvilleidae is provided.

Introduction

The symbiotic annelid, Eteonopsis geryonicola Esmark, 1874 (Polychae-
ta, Dorvilleidae), lives unattached among the gills and in the branchial cham-
ber of the deep-sea crab Geryon tridens Krgyer (Esmark, 1874; Bidenkap,
1895; Wesenburg-Lund, 1938; Desportes et al., 1977) and Geryon quinque-
dens Smith collected from the Atlantic Canyon (250—400 fm) in the western
North Atlantic. Wesenburg-Lund (1938) reported small worms most often
occurred in newly molted host crabs (G. tridens) and larger worms in crabs
which had not recently molted. Little is known about the life history of this
worm (e.g. entry into the host, mode of nutrition, reproduction, and survival
through host ecdysis).

A great deal of confusion exists in the literature concerning the original
description of Eteonopsis geryonicola. The following historical summary is
provided to alleviate such problems.

Eteonopsis geryonicola was named by Esmark (1874, often misquoted as
1873) in a brief summary of an oral presentation. A number of authors,
including Hartman (1951), Desportes et al. (1977), and Hartmann-Schroder
(1971) incorrectly date the description to 1878, presumably in reference to
a second oral presentation given by Esmark that year, but first published in
1880. Through a /Japsus the name was incorrectly given as E. geryonis in
the latter report. A more complete description of Esmark’s material fol-

VOLUME 94, NUMBER 1 Ta

lowed several years later (Bidenkap, 1895). Since the species is superficially
similar to the phyllodocid polychaete, Eteone, Bidenkap established a new
family for Eteonopsis: Pseudophyllodocidae. All specimens collected at that
time were from the branchial chambers of the red crab, Geryon; hence the
species name, E. geryonicola.

The generic name survived until Wesenburg-Lund (1938) referred Eteo-
nopsis to the eunicid genus Ophryotrocha, and submerged the family name,
Pseudophyllodocidae. Wesenburg-Lund (1938) must have been unaware
that Esmark provided the original description (albeit poor) and incorrectly
cited Bidenkap as the original author: Ophryotrocha geryonicola (Biden-
kap). Subsequent authors have recognized Esmark as the original author,
and referred the species to Ophryotrocha geryonicola (Esmark), now in the
Dorvilleidae. Until recently, this species has been known only from the
branchial chambers of two crabs, Geryon tridens Krgyer and G. quinque-
dens Smith. All other species of Ophryotrocha are free-living, interstitial
burrowers (Akesson, 1976).

In 1970, Fauchald established the family Iphitimidae for a number of
polychaetes found primarily in crustacean branchial chambers. Though
closely allied with dorvilleids, iphitimids could be distinguished in external
morphology by the presence of conspicuous branchiae across the dorsum.
Ophryotrocha geryonicola is abranchiate, as were all dorvilleids when Fau-
chald erected the Iphitimidae. Except for the presence of branchiae the
general external morphology of iphitimids and dorvilleids is similar. Several
analogous structures exist among the jaws of the two families as well. These
are discussed below.

Recent discovery of branchiate dorvilleid species further eliminated the
distinction between Iphitimidae and Dorvilleidae (Rullier, 1974; Armstrong
and Jumars, 1978). It is proposed, therefore, that the two families be com-
bined under Dorvilleidae, and furthermore, that the genus Eteonopsis Es-
mark, 1874 be revived for E. geryonicola to distinguish it from free-living
species of Ophryotrocha. Since the original description of E. geryonicola
was brief, a more complete description is provided herein.

This paper also reports on the occurrence of E. geryonicola in the red
crab, Geryon quinquedens off the U.S. east coast, as well as the first doc-
umented evidence of Eteonopsis inhabiting the gills and branchial cavity of
the Jonah crab, Cancer borealis. Collections were made at Station J-1 (Fig.
1) on the western Atlantic continental slope in May and August, 1977.

Occurrence Along United States East Coast

A total of 34 of 277 (12.3%) Geryon quinquedens and 1 of 197 (0.51%)
Cancer borealis contained polychaetes in the branchial chambers (Table 1).
In contrast, Wesenburg-Lund (1938) recorded infestation by this polychaete

78 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ATLANTIC
CITY

Fig. 1. Collection sites for host crabs, Geryon quinquedens and Cancer borealis. Virginia
Institute of Marine Science, Bureau of Land Management Outer Continental Shelf benchmark
study, 1975-1977. (Depths in meters.)

in 33% of 65 Geryon tridens examined in the Atlantic off Europe. Desportes
et al. (1977) reported it from the western Mediterranean. G. quinquedens
is quite common on the continental slope of the western North Atlantic
(Williams and Wigley, 1977) while the distribution and abundance of E.
borealis varies considerably with temperature (latitude) and depth. C. bo-
realis is a common inhabitant of the waters off the United States east coast
(MacKay, 1943) and is found in waters ranging in depth from several meters
to 750 meters (Squires, 1966). Of the specimens of C. borealis collected on
the continental shelf (<200 m), none was found to be infested with E. ger-
yonicola. C. borealis occurred rarely in slope samples and only one speci-
men contained the symbiotic worm. G. guinquedens, however, was taken
in large numbers from slope station J-1 (Fig. 1) and contained considerable
numbers (range 0-12) of E. geryonicola.

In agreement with Wigley et al. (1975), a substantial difference in the
occurrence of female vs. male G. quinquedens (Table 1) was found in the
present study, where males constituted only 9.7% of the sample. The per-
centage of infestation of worms in male hosts was significantly greater than

VOLUME 94, NUMBER | 79

Table 1.—Incidence of infestation by size and sex for the red crab, Geryon quinquedens by
the polychaete, Eteonopsis geryonicola.

Female Male

No. crabs collected 250 Bil
No. and (percent) infested 22 (8.8) 12, (44°74)
No. worms in infested crabs

mean and (range) 2M l= 7) 3.4 (1-12)
Carapace width—mean and (range)

of infested crabs (mm) 108.8 (87-122) 138.1 (122-151)
Carapace width—mean and (range)

of non-infested crabs (mm) 105.2 (84-122) 135.3 (120-144)

in females (chi-square test; a < 0.05). Intensity of infestation (number of
worms per host) was significantly greater for males than females (a < 0.05,
median test, Brown and Mood, 1951). The mean carapace width of infested
males (138.1 mm) was noticeably larger than that of the infested females
(108.8 mm) but no correlation between the occurrence of E. geryonicola
and the size of the crabs could be determined. Factors which govern the
investation of G. guinquedens and C. borealis by this dorvilleid cannot be
resolved without additional data.

Gut Analysis

A microscopic analysis of the gut contents of these worms revealed an
amorphous and acellular substance along with distinct clumps of bacteria.
This substance stained well with astrablue (specific for mucopolysaccarides)
and may, therefore, be digested mucus from the host crab. The question as
to whether these worms are commensals or parasites with a metabolic de-
pendence on the host, however, remains unclear.

Family Dorvilleidae Chamberlin, 1919
Eteonopsis geryonicola Esmark, 1874

Eteonopsis geryonicola Esmark, 1874:497—498.

Eteonopsis geryonis Esmark, 1880:14.

Eteonopsis geryonicola n. gen., n. sp. (Esmark, 1873) O. Bidenkap,
1895:72-74, pl. 3, figs. 1-3.

Ophryotrocha geryonicola (Bidenkap).—E. Wesenberg-Lund, 1938:1-14,
figs. 1-11.—Pettibone, 1961:181.—Hartman-Schroder, 1971:264.

Material examined.—NORWAY: Skagerrak, 4 specimens from branchial
chamber of Geryon tridens, B. Akesson, coll.—EAST COAST NORTH

80

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

0.5mm

0.02mm

VOLUME 94, NUMBER 1 81

&

Fig. 3. Eteonopsis geryonicola: A, Maxillae and mandibles of a juvenile (5S mm long) spec-
imen; B, Maxillae and mandibles of an adult (140 mm long) specimen; C, Mandibles of same.
(A, B, C after Wesenberg-Lund, 1938.)

AMERICA: 3 specimens from branchial chamber of Geryon quinquedens,
38°44'’N, 73°00'W, 350 m, D. Benner, coll. (USNM 55174). 2 specimens
from branchial chamber of Cancer borealis, 38°40.8'N, 73°04.3'W, 342 m,
D. Benner, coll. (USNM 54957). 5 specimens from branchial chamber of a
red crab, Atlantis Canyon, 39°50'N, 70°10’W, 250-400 fm, K. A. Wilhelm,
coll. (USNM 54339). 2 specimens from red crab, Gloucester, Maine, K. A.
Wilhelm, coll. (USNM 54338). 10 specimens from Geryon quinquidens,
38°44'N, 73°00’ W, 350 m, D. Benner, coll. (VIMS Acc. No. 736). 1 specimen
from Geryon quinquidens, 38°45.3'N, 73°01.0'W, 350 m, D. Benner, coll.
(VIMS Acc. No. 1250).

Description.—Specimens of Eteonopsis geryonicola range from 5—140

<—

Fig. 2. Eteonopsis geryonicola: A, B, Anterior and posterior ends in dorsal view of 100
mm specimen; C, Transverse section, anterior view of segment 73 with presetal lobe (pl) and
dorsal cirrus (dc) indicated; D, Simple falciger from middle notopodium; E, Composite falciger
from middle neuropodium; F, Simple falciger from same.

82 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

O.lmm

Fig. 4. Eteonopsis geryonicola: Maxillae and mandibles of an adult (100 mm long) speci-
men.

VOLUME 94, NUMBER | 83

mm in length, 0.3-5.0 mm wide, and 25 to several hundreds of segments
long. Body white in fresh and preserved specimens. Prostomium rounded
lobe with a pair of papilliform antennae and similar pair of palps (Fig. 2A).
No eyes present. Body gradually tapers posteriorly. Two short anal cirri
present (Fig. 2B). Between prostomium and first setigerous segment are 2
apodous and achaetous segments.

Maxillary carriers (Fig. 3B) fused medially, free ends form pincer-like
structure. Maxillae vary in number depending on size. Two small specimens
(40-50 mm) have 2 sets of maxillae with up to 14 paired maxillae total (Fig.
3A). Largest specimens (120—140 mm) have as few as 3 small paired maxillae
with Maxillae I (Mx I) fused to maxillary carriers (Fig. 3B). Intermediate-
sized specimens (ca. 100 mm) have large maxillary carriers fused to Mx I
and 6 additional maxillae (Fig. 4). Maxillary carriers and Mx I of similar
size and general form in all specimens 80-140 mm long. These structures of
smaller specimens poorly developed (Fig. 3A). Mandibles fused medially
(Figs. 3C, 4).

Parapodia uniramous and similar throughout. Conspicuous presetal lobe,
2 rounded postsetal lobes, and blunt dorsal cirrus present (Fig. 2C). A trans-
lucent aciculum bisects each parapodium. Each parapodium contains: dor-
sally—4-—8 long, simple, falcigers with blunt tip (Fig. 2D); ventrally—S-8
long, composite falcigers and 0—3 simple, hooked setae of similar length but
more narrow shafts (Fig. 2E, F). Branchiae lacking.

Remarks.—Bidenkap (1895) failed to mention the simple setae in the low-
er parapodial bundle. These are not present in all parapodia and may be
difficult to see. The most conspicuous difference in the morphology of
Ophryotrocha (Fig. 5) and Eteonopsis (Fig. 2) is the greater size of Eteo-
nopsis. Most species of Ophryotrocha are less than 10 mm long while Eteo-
nopsis may attain a length of 140 mm. In addition, Ophryotrocha species
are free-living, interstitial forms and E. geryonicola is known only as a
symbiont in crab branchial chambers. Ciliary rings present in many Ophry-
otrocha are lacking in Eteonopsis. The setae of the two genera differ. Blades
of composite falcigers are shorter in Eteonopsis (Figs. 2D, E, F; 5B, C).
Both genera, however, reportedly have a reduction in maxillae with age
(Wesenberg-Lund, 1938).

There are a number of similarities between the species of Iphitimidae
(Fig. 6) and Eteonopsis geryonicola (Fig. 2): both inhabit crab branchial
chambers; the mandibles of each (Figs. 3C, 7) are simplified and maxillary
carriers are fused with Mx I into pincer-like structures (Figs. 4, 7); setae of
both (Figs. 2D, E, F; 6C) include composite falcigers and simple hooks
(Hartnoll, 1962); both [phitime and Eteonopsis may occur in crab branchial
chambers, but certain species of [phitime (e.g. I. hartmanae) may also
occur beneath the abdomens of egg-bearing female crabs (Kirkegaard, 1977).

84 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Cc

Fig. 5. A, Ophryotrocha puerilis siberti, dorsal view of anterior end; B, simple and com-
posite falcigers; C, Ophryotrocha gracilis, simple and composite falcigers. (A, B, C after
Hartman-Schroder, 1971.)

Cc

Fig. 6. A, Iphitime loxorhynchi, anterior end in dorsal view, branchiae (b) indicated (after
Fauchald, 1977); B, J. hartmanae Kirkegaard, transverse section of middle segment with bran-
chiae (b) and dorsal cirri (dc) indicated (after Kirkegaard, 1977); C, I. loxorhynchi, simple and
composite falcigers (after Hartman, 1952).

VOLUME 94, NUMBER 1 85

Fig. 7. Iphitime loxorhynchi: Maxillae and mandibles.

The fusion of the maxillary carriers with Mx I in adult [phitime (Fauchald,
1970) and Eteonopsis distinguishes them from other Eunicea such as Onu-
phidae, Lumbrineridae, and Eunicidae whose pharangeal apparatus may
otherwise be similar. Both [phitime and Eteonopsis, like many dorvilleids,
have reduced maxillae in adult specimens (Fig. 3A, B). Eteonopsis may be
separated from [phitime by its lack of branchiae and presence of palps (Figs.
2A, C; 6A, B).

Key to the Genera of the Dorvilleidae
(after Jumars 1974)

1. Parapodia uniramous, without notoacicula ..................000- 4
— Parapodia sub-biramous, with enclosed notoaciculum ............ 2

86 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

2.. FURCALE;SEtae PRESENT. . Re oo ee 3
Furcate setae ‘absent: (0. .ee4 5 3 eee Dorvillea
3. Antennae small, simple; jaw pieces with 14 denticle rows.........
Win RARE BOR EES Sea Met coos Reiss See NA Ue. kata RO Ran eee Pettiboneia
— Antennae long, multiarticulate; jaw pieces with 4 denticle
TOWS rise do seapapeher so ansutnieutioiae 528 Rr ee Schistomeringos
4. vFurcate on geniculate setae present...) 4 ia eee 2)
Furcare and geniculate Setac.absentea.. eee eae 6
5. Palps well developed, with at least one article; jaws with 4 denticle
VOWS o0¥:.c'sy cetera ies ihe cman eer Sak en ae ee ne Protodorvillea
— Palps very small, simple or biarticulate; jaws with 2 denticle
TOW. Sere case ig ig 2rd interes eh UAE Tene eee ae Meiodorvillea
Gan Setaczallisimplesaciculan a...) ane. (eee ee Paraophyrotrocha
=, Setaeimelude sometfalcisers 4055. ©. 465 oe eee 7
7. Setiger 1 modified, with 2 types of large specialized setae: a simple
recurved spine and a large, thick, sharply recurved composite fal-
Cigenr, antennae and palpsi bianticulateme nee eke Exallopus
— Setiger | not modified; antennae and palps reduced, papilliform... 8
Sa Branchiae presentachoOssuGOnrsuinu en qe eee eee Iphitime
SruBranchiae: absent. 2320s used os bse gee a sn eee eee 9
9. Composite falcigers with elongate blades (Fig. 4B, C); body small
(Otmim) ainterstitialabunrowens) 4504 ane en eee Ophryotrocha
— Composite falcigers with short blades (Fig. 2E); body large (25—140
mm); symbiotic in crab branchial chamber ............... Eteonopsis
Acknowledgments

The authors are particularly grateful to Dr. Kristian Fauchald, Dr. Marian
Pettibone, and Dr. Meredith Jones of the United States National Museum
of Natural History for advice, constructive comments, and loans of museum
material. Additionally, Dr. Fauchald translated material concerning the
original description of the species and provided a critical review of this
manuscript. Dr. Bertil Akesson of the University of Gothenburg, Sweden,
kindly loaned specimens and provided suggestions on the revision of Eteo-
nopsis. The following Virginia Institute of Marine Science personnel are
gratefully acknowledged: Mr. Don Weston for critical review of this paper;
Dr. Frank O. Perkins for examination of bacteria from Eteonopsis digestive
tracts; Mr. Edward Matheson for help in collecting and handling of speci-
mens; Dr. Craig Ruddell for guidance in initiating this paper. This study
was supported in part by Bureau of Land Management contract number
AAS550-CT6-62.

Contribution no. 988 from the Virginia Institute of Marine Science.

VOLUME 94, NUMBER 1 87

Literature Cited

Akesson, B. 1976. Morphology and life cycle of Ophryotrocha diadema, a new polychaete
species from California.—Ophelia 15(1):23-35.

Armstrong, J. W., and P. A. Jumars. 1978. Branchiate Dorvilleidae (Polychaeta) from the
North Pacific.—Bull. South. Calif. Acad. Sci. 77(3):133-138.

Bidenkap, O. 1895. Systematisk Oversigt over Norges Annulata Polychaeta.—Vidensk. Selsk.
Christiania Forh. Volume for 1894. No. 10:72-74.

Brown, G. W., and A. M. Mood. 1951. On median tests for linear hypotheses.—Proc. 2nd
Berkeley Symp. Math. Statist. and Prob. (Univ. Calif. Press). 159 pp.

Desportes, I., L. Laubier, and J. Theodorides. 1977. Présence d’Ophyrotrocha geryonicola
(Esmark) (Polychete: Dorvilleidae) en Mediterranée Occidentale.—Vie Milieu 27(1):131-
133)

Esmark, L. 1874. Eteonopsis geryonicola.—Vidensk. Selsk. Christiania Forh. Volume for

1873:497-498.

. 1880. Eteonopsis geryonis.—Vidensk. Selsk. Christiania Forh. Volume for 1879:14.

Fauchald, K. 1970. Polychaetous annelids of the families Eunicidae, Lumbrineridae, Iphitim-

idae, Arabellidae, Lysaretidae and Dorvilleidae from western Mexico.—Allan Hancock

Monogr. Mar. Biol. 5:1—335.

. 1977. The polychaete worms, definitions and keys to the orders, families and genera.—

Nat. Hist. Mus. Los Angeles County, Science Series 28:1—190.

Hartman, O. 1951. Literature of the polychaetous annelids.—Los Angeles [privately printed].

290 pp.

. 1952. Iphitime and Ceratocephala (polychaetous annelids) from California.—Bull.

South. Calif. Acad. Sci. 51:9-20.

Hartman-Schroder, G. 1971. Annelida, Borstenwurmer, Polychaeta.—Tierwelt Dtl. 58:1—594.

Hartnoll, R. G. 1962. Iphitime cuenoti Fauvel (Eunicidae), a polychaete new to British
waters.—Ann. Mag. Nat. Hist. series 13, 5:93—96.

Jumars, P. A. 1974. A generic revision of the Dorvilleidae (Polychaeta), with six new species
from the deep North Pacific.—Zool. J. Linn. Soc. 54:101-135.

Kirkegaard, J. B. 1977. A new species of [phitime (Polychaeta: Iphitimidae) living under the
tail of Hyas (Crustacea: Decapoda) in the Oslo Fjord.—Allan Hancock Foundation
Special Publication: 199-207.

MacKay, O. C. 1943. Temperature and the world distribution of crabs of the genus Cancer.—
Ecology 24:113-115.

Rullier, F. 1974. Quelques Annélides Polychétes de Cuba recueillies dans les Eponges.—
Trans. Mus. Histoire Nat. ““Gr. Antipa’’ 14:9-77.

Squires, H. J. 1966. Serial atlas of the marine environment. Folio 12. Distribution of decapod
crustacea in the Northwest Atlantic.—Amer. Geogr. Soc.

Wesenburg-Lund, E. 1938. Ophryotrocha geryonicola (Bidenkap) (=Eteonopsis geryonicola
Bidenkap) refound and redescribed.—G6teborgs K. Vetensk. Handl., Ser. B. 6(8):1—13.

Wigley, R. L., R. B. Theroux, and H. E. Murray. 1975. Deep-sea red crab Geryon quinque-
dens survey off N.E. United States.—Mar. Fish. Rev. 37(8):1-21.

Williams, A. B., and R. E. Wigley. 1977. Distribution of decapod crustacea off northeastern
United States based on specimens at the Northeast Fisheries Center, Woods Hole,
Mass.—NOAA Tech. Rep. NMFS Circ. 407, 44 pp.

(GRG) Department of Biology, McNeese State University, Lake Charles,
Louisiana 70609; (DAB) Naval Polar Oceanography Center, Suitland Road,
F.O.B. No. 4, Suitland, Maryland 20023.

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 88-93

A NEW SPECIES OF FRESHWATER CRAB
(CRUSTACEA: ANOMURA: AEGLIDAE)
FROM INSULAR SOUTH CHILE

Carlos G. Jara and Maria T. Lopez

Abstract.—A new species, Aegla alacalufi (Crustacea, Anomura, Aegli-
dae) from Madre de Dios Island in far southern Chile is described. Mor-
phologically it closely resembles A. papudo Schmitt and A. concepcionensis
Schmitt from Chile and A. serrana Buckup and Rossi from Brasil. Its most
distinctive feature is the undivided telson plate, a character that is also
present in A. papudo with which the new species probably shares a common
ancestry.

Little information concerning the presence of members of the genus Aegla
in the insular and continental Chilean territory south of Puerto Montt is
available. Two species have been recorded from Chiloé Island (Haig, 1955;
Bahamonde and Lopez, 1963) and one from the Taitao Peninsula (46°30’S,
74°30’W) (Porter, 1917). The new species described here was collected on
Madre de Dios Island during cruise 72-2 of the oceanographic vessel Hero
to far southern Chile.

Aegla alacalufi, new species
Fig. 1

Holotype.—Instituto de Zoologia, Univ. Austral de Chile, IZUA C-471-
1, adult male collected in a brook at Puerto Henry, Madre de Dios Island
(50°01'10”S, 75°18’45”W), Ultima Esperanza, Chile, at sea level, on 7 Oc-
tober 1972 by H. Moyano.

Allotype.—IZUA C-471-2, adult female. Paratypes: IZUA C-471-3 and
MUZUC 16352 (Museo Zoologico, Univ. de Concepcion, Chile), 2 males;
IZUA C-471-4, C-471-5, MUZUC 16353, 4 females. Same locality and date
as holotype.

Diagnosis. —(1) Rostrum short and triangular with apical chitinous scale
amidst short hairs; (2) orbital spine absent; (3) rostral carina without scales;
(4) anterolateral angle of second abdominal epimeron rounded; (5) sternum
of fourth thoracic segment with medial, flattened, subdisciform offset; (6)
telson plate undivided.

Description of holotype.—Carapace longer than wide, ovoid; gastric area
conspicuously convex, appearing swollen and protuberant; dorsal surface

VOLUME 94, NUMBER 1 89

of carapace and chelae quite punctate and sparsely studded with fine bris-
tles. Chitinous scales limited to those areas mentioned below.

Rostrum short, triangular, acuminate, with broad base and depressed dor-
sally; apical scale concealed by bristles; rostral carina moderately well de-
veloped, rounded dorsally, decreasing in height anteriorly and terminating
near rostral tip; surface smooth but somewhat hairy; flanking troughs lack-
ing.

Orbits wide; orbital spine absent although weak tubercle present on lateral
margin; anterolateral angles of carapace short and blunt. Epigastric areas
semicircular, protuberant, punctate, and sparsely setiferous; protogastric
eminences inconspicuous. First hepatic lobe with margin slightly upturned,
limited frontally by wide, shallow notch; second and third hepatic lobes
poorly delineated.

Anterolateral branchial lobe short and blunt, edge of branchial areas hairy
and somewhat nodose; cardiac area wide and trapezoidal; areola subrectan-
gular and moderately convex. Dorsum of abdominal terga strongly convex;
anterolateral angle of second epimeron rounded and hairy, its ventrolateral
edge straight; telson plate undivided, its lateral margins straight and apex
rounded. Sternum of fourth thoracic segment with broad low anteromedian
tubercle, free border of which subdisciform, border immediately lateral to
tubercle somewhat concave, and lateral margin of sternum elevated ven-
trally.

Antennal flagellum long and slender, longer antenna 1.25 times as long as
cephalothorax. Chelipeds slightly robust, equal in size; chelae subrectan-
gular and not inflated, their dorsal and ventral surfaces densely punctate
and covered with fine setae, latter longer on dactylus, palmar crest, and
ventrolateral margin of fixed finger; blunt nodule present near dorsal base
of dactyl, that on left chela with minute apical scale; palmar crest insignif-
icant, not surpassing contour of chela, its free border nodulated; dorsum of
carpus scabrous with arcuate row of low blunt tubercles flanked apically by
transversely aligned hairs; carpal crest clearly defined, nodulate proximal
to long, conical, slightly curved distal spine, latter separated from anterior
carpal lobe by wide semicircular notch; apex of carpal lobe blunt and in-
clined toward distal end of carpus; ventral margin of merus nodulate, dorsal
margin with longitudinal row of 9 or 10 small blunt tubercles; distodorsal
border smooth and somewhat hairy; ventral margin of ischium with low
non-acute swelling at proximal end; remaining pereiopods long and slender;
dactylus of fourth pereiopod longer than that of second.

Description of allotype.—Differing from holotype in following respects:
rostral carina prominent and well marked almost to rostral apex; middorsal
line on gastric area slightly elevated, darker, and covered by narrow band
of short setae; anterior margin of protogastric eminences well delimited and
bearing sparsely and irregularly placed hairs; proximal nodules of palmar

90 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Aegla alacalufi, n. sp. (all illustrations from holotype except 1 from female P:): a,
Dorsal view; b, Carpus of left cheliped; c, Telson plate; d, Third and fourth thoracic sterna;
e, Dorsal view of rostral area; f, Lateral view of anterior part of cephalothorax; g, Inner ventral
margin of ischium of left cheliped; h, Lateral view of second abdominal epimeron; i, Telson
plate.

crest inconspicuous; apex of telson plate slightly notched and devoid of
plumose setae.

Color.—(Alcohol fixed.) Gastric and cardiac surfaces bluish white (Cy-
aneus 1), other dorsal cephalothoracic and abdominal surfaces yellowish
brown tan (Flavus 3); pereiopods smoky tan (Flavus 5); dorsal surface of
carpus and propodus of cheliped yellowish white, somewhat irridescent;
ventral surfaces marmoraceous to light brown or ferrugineous. (Color stan-
dards according to Paclt, 1958.)

Variations and measurements.—The type-series is morphologically ho-
mogeneous, the shape of the telson constituting the most variable character.
All specimens have undivided telsons without a trace of a longitudinal su-
ture; however, | male and 3 females have an obvious semicircular apical
notch of variable depth, the margin of which lacks plumose setae. The telson
with the deepest notch is depicted in Fig. li. Less conspicuous is the vari-
ation in the shape of the rostrum which varies from distinctly triangular to
slightly ligulate, the apical part broadened and flattened. The chelae of the
female are less robust than in the male, but the difference is not remarkable.

VOLUME 94, NUMBER 1 9]

Table 1.—Somatometry of A. alacalufi n. sp., type-series. All measurements in mm; M:
male; F: female; Holo: holotype; Allo: allotype; P, to P,: paratypes.

IZUA Ce IZUACCan MIZUNO F EMUZUC MIZUAG SeMUZUC = IZUA-C = MUZUC
471-

Collection 471-1 471-2 1-3 16352 471-4 16353 471-5 16353
Specimen: Holo Allo Pi PS Jee P, Pe JP,
Sex: M F M M F F F F
GE: 10.6 9.5 10.2 9.4 10.2 8.6 8.5 8.5
RL: ey), 2.0 Dell 2.0 Df 1.9 1.9 1.8
IRCILS Teel: 6.9 18 6.8 V3 6.2 6.1 6.1
FW: 355) 3.2 3.4 367) 3.4 3.0 3.0 2.8
PCW: 6.6 6.0 6.3 5.8 6.2 5533 Se Dall
CW: 8.2 7.8 8.1 ee 8.1 6.8 7.0 6.5
ECE: Weil 11.0 — 10.7 12.0 10.1 9.9 9.4
RCL: 12.4 11.0 11.8 10.7 12.0 10.0 9.8 9.4
L2PL: IY 12.4 13.8 11.8 Be) 11.4 MEG 10.4
L2DL: 3.8 Sead 3.4 2 all 3.4 2.9 2.9 Dell
L4DL: 4.0 3.6 3.8 3.4 3.9 357) sal 3.0

Wee Dodh Zell Me 1\ 2.0 hp 1.9 169 1.9

Measurements of the specimens constituting the type-series are presented
in Table 1. These were made with the aid of calipers to the nearest 0.1 or
0.2 mm, depending on the size of the structure. Areas or structures mea-
sured are defined as follows: CL, carapace length, distance between rostral
apex and posterior margin of cephalothorax; RL, rostral length, distance
between rostral tip and midpoint of transverse line tangent to deepest points
of orbital margins; PCL, precervical length, distance between rostral tip and
midpoint of cervical groove; FW, frontal width, distance between tips of
anterolateral angles of carapace; PCW, maximum precervical width, dis-
tance across third hepatic lobes; CW, maximum carapace width; LCL, left
cheliped length; RCL, right cheliped length; L2PL, length of second left
pereiopod; L2DL, dactylar length of second left pereiopod; L4DL, dactylar
length of fourth left pereiopod; TL, telson length (when apical notch present,
posterior margin considered as line joining caudalmost parts of telson).

Natural history.—Little information is available; the type-series was col-
lected on the sandy bottom of a forest brook outlet to the sea. Some spec-
imens were found under detached Macrocystis Agardh (Phaeophyta) hold-
fasts about 50 m from the seashore.

Distribution.—Known only from the type-locality.

Etymology.—Latin genitive singular form of Alacalufes, a tribe of aborig-
ines that inhabited the southern Chilean Archipelago (Encyclopaedia Bri-
tannica, 1961).

Comparison.—Aegla alacalufi should be included in the *‘Pacific rostrum
type’ group of species proposed by Schmitt (1942). Morphologically it re-
sembles A. concepcionensis Schmitt (1940) and A. papudo Schmitt (1942)

92 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

from Chile and A. serrana Buckup and Rossi (1977) from Brasil. It shares
the following features with them: dorsum of gastric area swollen and strong-
ly convex; areola wide, subrectangular, and slightly convex; orbital spine
absent, and apex of carpal lobe displaced distally. It differs from these
aeglids as follows: rostral carina smooth and lacking scales; propodus of
chela not inflated; palmar crest reduced, its free edge nodulated; upper
longitudinal margin of merus of cheliped without spines; and sternum of
fourth thoracic segment with anteromedial prominence. Aegla alacalufi re-
sembles A. concepcionensis and A. serrana in lacking well defined proto-
gastric eminences, and the inner ventral margin of the ischium of the che-
liped bears neither spines nor scales. As in A. papudo and A. serrana, the
anterolateral angle of the second abdominal epimeron of A. alacalufi is
rounded. The latter differs from A. concepcionensis and A. papudo in lack-
ing a small concavity behind the dorsal base of the anterolateral angle of
the carapace. A comparison of the type-series of A. alacalufi with a sample
of A. papudo (IZUA C-414: 1 male, 6 females, adults) from La Ligua
(Aconcagua, Chile) revealed that both species have an undivided telson
plate. Dr. Alan Solem (Field Museum of Natural History, Chicago, U.S.A.)
(personal communication) confirmed that this character is also present in
the type-series of A. papudo. None of the specimens of A. papudo exam-
ined, however, has a notch on the apex of the telson.

Remarks.—The finding of A. alacalufi at Madre de Dios Island extends
the previously known range of the Aeglidae about 400 km to the south (see
Porter, 1917; and Ringuelet, 1948).

Telson morphology places A. alacalufi and A. papudo in a subgroup apart
from the other members of the genus Aeg/a in which it is dimerous. It is
difficult at the present, however, to judge the systematic value of the char-
acter, particularly its relevance in establishing a new generic or subgeneric
category. Unfortunately, previous authors have not consistently noted
whether or not the telson of the species described by them was divided.
Leach, 1821, and Hobbs, Hobbs, and Daniel, 1977, included the longitudi-
nally articulated telson in their diagnoses of the genus, unaware of the un-
divided plate in A. papudo. Whereas the structure of the telson suggests a
common ancestry for A. alacalufi and A. papudo, the possibility of mor-
phological convergence in disjunct lineages cannot at present be dis-
regarded.

Acknowledgments

The authors thank Prof. Hugo Moyano (Universidad de Concepcion,
Chile) for presenting to them the valuable sample of A. alacalufi, Dr. Alan
Solem (Field Museum of Natural History, Chicago, U.S.A.) for examining
the telsons of the type-series of A. papudo, Dr. H. H. Hobbs, Jr. (Smith-

VOLUME 94, NUMBER | 93

sonian Institution, Washington, U.S.A.) for his thorough revision of the
manuscript, and Mrs. Patricia Jara for her assistance in preparing the text
in English.

Literature Cited

Bahamonde, N., and M. T. Lopez. 1963. Decapodos de aguas continentales en Chile.—Inv.
Zool. Chilenas 10:123-149.

Buckup, L., and A. Rossi. 1977. O Genero Aegla no Rio Grande do Sul, Brasil (Crustacea,
Decapoda, Anomura, Aeglidae).—Rev. Brasil. Biol. 37(4):879-892.

Haig, J. 1955. The Crustacea Anomura of Chile.—Rep. Lund Univ. Chil. Exped. 1948-49,
20, Lund Univ. Arsskrift. N.F. Avd. 2, Bd 51, Nr 12, 68 pp.

Hobbs, H. H., Jr., H. H. Hobbs III, and M. A. Daniel. 1977. A review of the troglobitic
decapod crustaceans of the Americas.—Smithsonian Contrib. Zool. 244:1-183.

Leach, W. E. 1821. Galateadées, pages 49-56 in F. G. Levrault, ed., Dictionnaire des Sciences
Naturelles, 18 (1820). Strasbourg.

Paclt, J. 1958. Farbenbestimmung in der Biologie.—Gustav Fisher Verlag, Jena.

Porter, C. 1917. Los crustaceos de la expedicion Taitao.—Bol. Mus. Nac. Hist. Nat., Chile
10:94-101.

Ringuelet, R. 1948. Los cangrejos argentinos del genero Aegla de Cuyo y la Patagonia.—Rev.
Mus. La Plata, nov. ser., Zool. 5:297—-349.

Schmitt, W. L. 1940. Two new species of Aegla from Chile.—Rev. Chilena Hist. Nat.

44(1940):25-31, pl. 1.

. 1942. The species of Aegla, endemic South American freshwater crustaceans.—Proc.

U.S. Nat. Mus. 91:431—-520.

(CGJ) Instituto de Zoologia, Universidad Austral de Chile, casilla 567,
Valdivia, Chile; (MTL) Instituto Central de Biologia, Universidad de Con-
cepcion, Chile.

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 94-100

TAXONOMIC OBSERVATIONS ON EASTERN
PACIFIC ANTITHAMNION SPECIES
(RHODOPHYTA: CERAMIACEAE)
DESCRIBED BY E. Y. DAWSON

David N. Young

Abstract.—Three of the six species of the red algal genus Antithamnion
described by E. Y. Dawson are studied critically. Re-examination of the
type specimens and study of new material of A. hubbsii, A. pseudocorti-
catum, and A. mcnabbii reveal that the latter two are more correctly trans-
ferred to Balliella and Antithamnionella respectively. Gland cells previ-
ously unrecognized for B. pseudocorticata are described, and tetrasporangia
reported for the first time for Antithamnionella mcnabbii and Antithamnion
hubbsii. Distribution and new records are noted for each taxon.

E. Yale Dawson (see Abbott, 1966) described a number of new taxa from
the eastern North Pacific, including 6 species of Antithamnion Nageli (1847)
(Ceramiaceae, Ceramiales) from the Gulf of California and the Pacific coasts
of California, Mexico, and Central America: A. breviramosus Dawson
(1949), A. dumontii Dawson (1960), A. hubbsii Dawson (1962), A. mcnabbii
Dawson (1950), A. pseudocorticatum Dawson (1962), and A. scrippsiana
Dawson (1949). Later, Dawson (1953) recognized A. scrippsiana to be a
form of A. glanduliferum Kylin (1925), which was later transferred by Wol-
laston (1971) to Antithamnionella Lyle (1922). She also considered Anti-
thamnion breviramosus Dawson in this genus, as Antithamnionella brevi-
ramosa (Dawson) Wollaston (in Womersley and Bailey, 1970). Abbott (1979)
transferred A. dumontii to Wrangelia C. Agardh (1828). The remaining
species of Dawson, Antithamnion hubbsii, A. mcnabbii, and A. pseudo-
corticatum, were known only from the type collections (all sterile and their
taxonomic status was incompletely known). New observations on the type
specimen and on additional material identified as these species add new
information on these species and, in the case of Antithamnion pseudocor-
ticatum and A. mcnabbii, justify their transfer to other genera.

Balliella pseudocorticata (Dawson) comb. nov.
Figs. 1, 2

Basionym.—Antithamnion pseudocorticatum Dawson, Allan Hancock
Pacific Expedition 26:20, Pl. 7, Figs. 1, 5S (1962).
Description.—Erect thallus branches to 5 mm high arising from prostrate

VOLUME 94, NUMBER 1 95

axes attached to substrate by long, multicellular rhizoids; branches of erect
axes in opposite pairs, slightly incurved, distichous, bearing opposite pairs
of 1-3 celled often once-branched, incurved whorl branchlets and with
smaller, subrectangular basal cell; lateral branches with opposite whorl
branchlets and with same branching as erect axes; whorl branches and
branchlets incurved but not surrounding apical cell and bearing whorl
branchlet initials in irregular sequence; basal cells of branches and whorl
branchlets each bearing two adaxial and one to two abaxial branchlets, these
remaining meristematic and becoming compound, enveloping basal portions
of major branches and forming loose cortication. Gland cells spherical, (8—)
11 (-13) wm diameter, born abaxially on basal cell of whorl branchlets
(Fig. 1). Gland cells usually occurring singly, but two gland cells may be
present on a single whorl branchlet basal cell in older portions of the thallus.
Corticating branchlets also with gland cells. Reproductive material not
found.

Holotype.—Dawson 6934b (AHFH 71122), 15.11.49, San Lorenzo Chan-
nel, dredged between Isla Espiritu Santo and Baja coasts, near La Paz, Baja
California del Sur, Mexico.

Additional records.—Galapagos Islands, R/V Searcher station 327, near
Punta Espinosa, Isla Fernandina (P. C. Silva, AHFH 85054).

Remarks.—Dawson (1962, p. 20—21) accurately describes the plant except
for the statement ‘‘gland cells absent,’’ which is in error. It is unclear why
Dawson failed to observe the gland cells in the type material. The distinc-
tive, refractive nature of certain types of living glandular cells is lost by
fixation in formalin (Young, 1977), but the cells on the formalin-preserved,
Karo-mounted type specimen are clearly present (Fig. 2).

The morphology of Antithamnion pseudocorticatum differs from the char-
acteristic vegetative features which define the genus Antithamnion (see
Wollaston, 1968): axes completely lacking rhizoidal cortication; pinnae dis-
tichous and either opposite, alternate, or secund and with a small basal cell
which does not bear pinnules; gland cells born on special short branches.
The Gulf of California plant agrees with the features of the genus Balliella
Itono and Tanaka (1973) in apical organization, branching pattern, devel-
opment of nodal cortication of the axes, and unique gland cell shape and
position. Balliella bears cruciately divided, adaxial tetrasporangia, and
many procarps on the basal cell of subapical lateral branches. Unfortu-
nately, the absence of reproductive material on A. pseudocorticatum pre-
cludes comparison of reproductive structures, but the vegetative features
alone are strong evidence warranting transfer of A. pseudocorticatum to
Balliella.

Prior to these observations, Balliella contained two species, B. crou-
anioides (Itono) Itono and Tanaka and B. subcorticata (Itono) Itono and
Tanaka. Balliella pseudocorticata is easily distinguished from B. crou-

96 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

—_ ie :
Figs. 1, 2. B. pseudocorticata: 1, Living specimen with adaxial gland cells; 2, Holotype
with gland cell. Figs. 3, 4. A. mcnabbii: 3, Habit; 4, Tetrasporangium. Fig. 5. A. Aubbsii,
pinnules with tetrasporangia. Figs. 1, 2, 4, 5, bar is 50 wm; Fig. 3, bar is 200 um.

fe

#2

VOLUME 94, NUMBER 1 97

anioides by its irregular branching pattern and the compact nature of the
nodal cortication in the latter species. Balliella pseudocorticata is smaller
and stouter than B. subcorticata, but otherwise seems similar. The recently
described Bakothamnion curassavicum van den Hoek (1978) from the east-
ern Caribbean may be placed more properly in Balliella (van den Hoek,
pers. comm.). In view of environmental effects on thallus form in the Ce-
ramiaceae (Murray and Dixon, 1973), re-examination of variation in B. sub-
corticata and comparisons with the Caribbean and western Pacific plants is
desirable.

Antithamnionella mcnabbii (Dawson) comb. nov.
Figs. 3, 4

Basionym.—Antithamnion mcnabbii Dawson, Los Angeles County Mu-
seum Contributions in Science 27:28, Fig. 7C. 1959.

Description.—Erect thallus minute, to 8 mm tall (Fig. 3), attached to
substrate by unbranched, multicellular rhizoids originating from basal cell
of lateral branchlets; axes ecorticate; whorl branchlets short, slightly in-
curved, usually in threes, forked 3-5 times with a digitate appearance, from
subrectangular basal cells; axial cell bearing lateral branches also with a
whorl of three branchlets; lateral branches 304 segments apart with whorl
in branchlets as in main axis; gland cells absent in type; tetrasporangia (Fig.
4) spherical to subspherical, tetrahedrally divided, sessile, frequently with
additional branchlets on the basal cell of a lateral branch; male and female
structures unknown.

Holotype.—Dawson 18855, 20.iv.58, from El Solitario Rock, Bahia Agua
Verde, Baja California del Sur, Mexico (LAM now AHFH 81902).

Additional records.—I have examined material of this species from the
Gulf of California, now deposited in AHFH and US. Sterile material from
Isla Fernandina, Galapagos Islands (P. C. Silva, AHFH 85053) and tetra-
sporic material from Caleta Santa Maria, Baja California del Sur (J. Norris
3406, US) was examined.

Remarks.—Tetrasporic material from the Gulf of California (J. Norris
3406) clearly exhibits stages in the simultaneous, oblique cleavage process
characteristic of tetrahedral division (Wollaston, 1968). The previous report
(Itono, 1969) of cruciate tetrasporangia in Antithamnion mcnabbii from Ja-
pan is considered by Itono (1977) to be a doubtful record for that species.
The Japanese specimen is similar to Mexican specimens, and Itono’s (1969)
Fig. 5A, if judged alone, could represent tetrahedral division.

This species exhibits features in common with Antithamnionella Lyle
(1922) and excluding it from Antithamnion Nageli (Wollaston, 1968): the
shape and tetrahedral cleavage of tetrasporangia, the pinnae in whorls of
three, the shape of basal cell of pinnae, the absence of gland cells on special
gland cell branches. Likewise, the branch morphology and the sessile tet-

98 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

rasporangia of A. mcnabbii exclude this species from Scagelia Wollaston
(1971) and Hollenbergia Wollaston (1971). The digitate morphology of the
whorl branchlets and absence of gland cells distinguish Antithamnionella
mcnabbii from the North Pacific species of this genus, A. glandulifera (Ky-
lin) Wollaston and A. pacifica (Harvey) Wollaston and from the Japanese
species, A. miharai (Tokida) Itono (1977). Vegetatively, A. mcnabbii seems
most closely related to A. breviramosa (Dawson) Wollaston, also from Pa-
cific Mexico and the Gulf of California (Dawson, 1962), which has consid-
erable variation in numbers of gland cells present.

Antithamnion hubbsii Dawson
Fig. 5

Antithamnion hubbsii Dawson, Allan Hancock Pacific Expeditions 26:16,
Ries) hice Pino hice Sa looo.

Description.—The vegetative aspects of this species were described com-
pletely by Dawson (1962) and Wollaston (in Abbott and Hollenberg, 1976).
Recent California collections identified as this species allow the following
description of previously unknown tetrasporangia. Mature tetrasporangia
(Fig. 5) are ovoid, 35 x 60 um, sessile, usually in place of gland cells on
pinnules of first or second cell (not basal cell) of pinnae on erect axes.

Holotype.—Dawson 8302 (AHFH 71130), 19.xii.49, from 70 m depth,
Melpomene Cove, Isla Guadalupe, Mexico (R/V Velero station 1919-49).

Additional records.—Various collections from subtidal areas at Santa
Catalina Island, California (Hageman 24, AHFH; Young 998), and from the
Dana Point Marina, Orange County, California (AHFH 85052, 85053) were
examined. Previously known only from type locality and Santa Catalina
Island (Wollaston in Abbott and Hollenberg, 1976).

Remarks.—The tetrasporangial and vegetative features of this distinc-
tive species are consistent with its placement in Antithamnion. Antitham-
nion hubbsii has similarly placed, but smaller, tetrasporangia than the sim-
ilar A. callocladus Itono (1971), which is overall more robust and contains
1-2 more orders of branching than the former species.

California specimens differ somewhat from the holotype in that the basal
cells of the pinnae on prostrate axes bear two attachment rhizoids whereas
single rhizoids are formed from pinnae on erect axes.

This species is relatively abundant in appropriate habitats in southern
California, but gametangial material remains unknown. To date, my at-
tempts to obtain gametophytes by cultivation of tetraspores have been un-
successful.

Acknowledments

I am grateful to Drs. P. C. Silva (Herbarium, University of California,
UC) and J. N. Norris (U.S. National Herbarium, Smithsonian Institution,

VOLUME 94, NUMBER I 99

US) for kindly providing specimens from their collections. R. Setzer, Drs.
M. J. Wynne and J. N. Norris made helpful comments on the manuscript,
and R. Moe provided useful discussions. This is contribution number 381
from the Allan Hancock Foundation.

Literature Cited

Abbott, I. A. 1966. Elmer Yale Dawson (1918—1966).—Journal of Phycology 2:129-132.

, and G. J. Hollenberg. 1976. Marine Algae of California.—Stanford University Press,

Stanford, California. 827 pp.

Agardh, C. A. 1828. Species Algarum Rite Cognitae, cum Synonymis, Differentiis Specificis
et Descriptionibus Succinctis, Vol. 2.—Mortiz, Greifswald, Germany. 189 pp.

Dawson, E. Y. 1949. Contributions toward a marine flora of the southern California Channel

Islands.—Allan Hancock Foundation Occasional Papers 8:1—57.

. 1954. Notes on Pacific Coast marine algae, VI.—Wasmann Journal of Biology 11:323-

35 1k

. 1959. Marine Algae from the 1958 cruise of the Stella Polaris in the Gulf of Califor-

nia.—Los Angeles County Museum Contributions in Science 27:1-39.

. 1960. New records of marine algae from Pacific Mexico and Central America.—Pacific

Naturalist 1:31—53.

. 1962. Marine red algae of Pacific Mexico. Part 7. Ceramiales: Ceramiaceae, Deles-

seriaceae.—Allan Hancock Pacific Expedition 26:1—207.

Hoek, C. van den. 1978. Marine algae from the coral reef of Curacao, Netherland Antilles,
I: Three new and one rarely observed species from the steep fore-reef.—Aquatic Botany
5:47—62.

Itono, H. 1969. The genus Antithamnion (Ceramiaceae) in southern Japan and adjacent

waters, I.—Memoirs of the Faculty of Fisheries, Kagoshima University 18:29—45.

. 1971. The genus Antithamnion (Ceramiaceae) in southern Japan and adjacent waters,

II].—Memoirs of the Faculty of Fisheries, Kagoshima University 20:209-216.

. 1977. Studies on the Ceramiaceous Algae (Rhodophyta) from Southern Parts of Ja-

pan.—Bibliotheca Phycologica Bd. 35. J. Cramer, Vaduz. 499 pp.

, and T. Tanaka. 1972. Balliella, a new genus of Ceramiaceae.—Botanical Magazine,

Tokyo 86:241-252.

Kylin, H. 1925. The marine red algae in the vicinity of the biological station at Friday Harbor,
Washington.—Lunds Universitets Arsskrift N.F. 21:1-87.

Lyle, L. 1922. Antithamnionella, a new genus of algae.—Journal of Botany 60:346—350.

Murray, S. N., and P. S. Dixon. 1975. Effect of light intensity and light period on the devel-
opment of thallus form in the marine red alga Pleonosporium squarrulosum (Harvey)
Abbott (Rhodophyta: Ceramiales), I: Apical cell division.—Journal of Experimental Ma-
rine Biology and Ecology 13:15-27.

Nageli, C. 1847. Die neuern Algensysteme und Versuch zur Begrundung eines einen Systems
der Algen und Florideen.—Neue Denkschriften Allgemeine Schweiz Gesellschaft Na-
turwissenschaftlich 9(2):1—275.

Wollaston, E. M. 1968. Morphology and taxonomy of southern Australian genera of Crouan-

ieae Schmitz (Ceramiaceae, Rhodophyta).—Australian Journal of Botany 16:217—417.

. 1971. Antithamnion and related genera occurring on the Pacific Coast of North Amer-

ica.—Syesis 4:73-92.

—. 1972. Generic features of Antithamnion (Ceramiaceae, Rhodophyta) in the Pacific
region. Pp. 142-145 in Nisizawa, K. et al. (editors), Proceeding 7th International Sea-
weed Symposium. Sapporo, Japan, 1971. J. Wiley & Sons, New York.

100 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Womersley, H. B. S., and A. Bailey. 1970. Marine Algae of the Solomon Islands.—Philo-
sophical Transactions Royal Society of London 259:275-352.

Young, D. N. 1977. Comparative Fine Structure and Histochemistry of Vesiculate Cells in
Selected Red Algae. Unpublished Ph.D. dissertation, University of California, Berkeley.
155 pp. (Dissertation Abstracts #7812834, University Microfilms, Ann Arbor, Mich.).

Allan Hancock Foundation and Department of Biological Sciences, Uni-
versity of Southern California, Los Angeles, California 90007.

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 101-106

A REVISION OF THE NORTHERN FORMS OF
EUPHONIA XANTHOGASTER (AVES: THRAUPIDAE)

Storrs L. Olson

Abstract.—Analysis of geographic variation in populations of Euphonia
xanthogaster from Panama, Colombia, and Venezuela, shows E. x. cho-
coensis to be divisible into a small lowland subspecies, E. x. chocoensis
Hellmayr, in western Colombia and Ecuador, and a larger upland one, E.
x. oressinoma, new subspecies, in the western and central Andes and on
the western slope of the northern part of the eastern Andes. E. x. badissima,
new subspecies, from the Sierra Perija south to the Tachira Valley, and
Boyaca, Colombia, is distinguished from E. x. exsul Berlepsch of northern
Venezuela. E. x. badissima and E. x. oressinoma intergrade in Santander
del Norte, the intergrades having previously been misidentified as E. x. bre-
virostris Bonaparte, which now appears to be unknown north of Cundina-
marca in the eastern Andes.

The following revision treats the subspecies of the Orange-bellied Eu-
phonia, Euphonia xanthogaster, that occur in Panama and most of Colom-
bia and Venezuela. I have considered only those populations that have in
the past been recognized under the names chocoensis, exsul, and breviros-
tris. The distribution of these taxa, two new subspecies, and their inter-
grades, is shown in Fig. |. The analysis was based mainly on adult male
specimens.

Euphonia xanthogaster chocoensis Hellmayr

Euphonia xanthogaster chocoensis Hellmayr 1911, Rev. Frang. d’Orn. 2:23.
Rio Cajon, Choco, Colombia.

Characters.—Small, wing of adult males 60 mm or less (55.8—60.8, av-
erage 58.6, n = 16), bill small. Cap yellowish, underparts yellow, only faint-
ly tinged with ochraceous.

Range.—Lowlands of western Colombia west of the Andes, from Choco
south through Narino into Ecuador at least as far as Esmeraldas and Im-
babura.

Specimens examined.—COLOMBIA. CHOCO: Bahia de Solano (1,
AMNH); Rio Uva mouth, near Rio Bojaya (1, AMNH); Rio Nuqui, base of
Baudo Mountains (6, USNM); Rio Jurubida, Baudo Mountains (3, USNM);
E] Tambo (6, CM); Potedo (2, CM); Quibdo, Rio Atrato (2, CM). VALLE:

102 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

vs

EUPHONIA XANTHOGASTER
CHOCOENSIS
ORESSINOMA
BREVIROSTRIS
BADISSIMA

EXSUL

erPP@eo

ORESSINOMA X BADISSIMA

Fig. 1. Outline map showing the distribution of the northern forms of Euphonia xantho-
gaster in Colombia, Venezuela, and eastern Panama, based on specimens examined in this
study. Fine lines indicate provincial boundaries; cross-hatched areas indicate elevations above
1000 m.

Malaguita (2, CM); Cordoba (2, CM; 2, ANSP); Punto Muchimbo, just be-
low mouth of Rio Calima (1, USNM). CAUCA: Novita, Rio San Juan (1,
USNM). NARINO: Barbacoas (2, AMNH). ECUADOR. ESMERALDAS:
Cachabi (4, AMNH). IMBABURA: Lita (1, AMNH).

Remarks.—This is a small form restricted to the lowlands west of the
western Andes. Two males from Rio Jurubida in the Baud6 Mountains have
the wing slightly longer than typical chocoensis (61.0, 61.7), but are never-
theless smaller than all but two specimens of the following subspecies.

Euphonia xanthogaster oressinoma, new subspecies

Holotype.—USNM 436904, adult male, Hacienda Sofia, Rio Samana, Cal-
das, Colombia, elevation 3750 feet (1143 m). Collected 26 May 1951 by M.
A. Carriker, Jr., original number 20308.

VOLUME 94, NUMBER 1 103

Characters.—Coloration like that of chocoensis but size larger, wing of
adult males usually 63 mm or longer (61.0—68.6, average 63.9, n = 35), bill
noticeably larger.

Range.—Panama in Darién, and in Colombia on both slopes of the west-
ern Andes south at least to Narino, and the western slope of the central
Andes; also known from a widely disjunct locality on the western slope of
the eastern Andes in Santander.

Specimens examined.—PANAMA. DARIEN: Rio Jaqué, mouth of Rio
Imamado (1, USNM); Rio Jaqué, Las Penitas (1, USNM); 9 mi E of Cana
(2, USNM); Cerro Pirre (4, USNM); Cerro Sapo (1, ANSP).

COLOMBIA. ANTIOQUIA: Hacienda Potreros, 15 mi (24 km) SW of
Frontino, Rio Herradura (4, USNM); Valdivia, above Sevilla (3, USNM);
La Bodega, N side of Rio Negrito on Sonson-Narino highway (1, USNM);
Alto Bonito (1, AMNH). VALLE: Km 18 on Cali-Buenaventura road (1,
USNM); Height of Caldas (5, CM); Bitaco Valley (2, CM); La Cumbre (1,
CM; 1, ANSP); San Antonio (1, AMNH). CALDAS: Hacienda Sofia, Rio
Samana (8, USNM); La Selva (4, ANSP). CAUCA: Cocal, W of Popayan
(1, AMNH); E of Palmira (2, AMNH); El Tambo (5, ANSP). NARINO:
Ricuarte (4, ANSP); Mayasquer (1, ANSP). SANTANDER: Virolin, 28 km
S of Charala (5, USNM).

Etymology.—Greek oressinomos, ranging the mountains.

Remarks.—Hellmayr (1936:26) has already noted that birds from the high-
land portions of the range attributed to chocoensis are larger than those
from the lowlands. Inasmuch as these are consistently separable, I have
chosen to recognize this variation nomenclaturally. Measurements do not
adequately convey the noticeably smaller size of chocoensis, particularly
of the bill. Caution must be used in taking wing measurements because adult
males in definitive plumage often retain primaries from the previous subadult
plumage, and these consistently give a smaller measurement. Such speci-
mens were eliminated from consideration in the measurements discussed
above.

There is some variation within oressinoma; birds from Panama tend to
be somewhat darker, more ochraceous below, and those from Caldas are
yellowest, but individual variation bridges the differences. The specimens
reported here from Virolin in Santander apparently constitute the first
record of “‘Chocoensis’’-like birds from the eastern Andes and are widely
separated from the nearest known population of oressinoma. Nevertheless,
I cannot distinguish them in any way from topotypes of oressinoma from
Caldas, although all the specimens from Santander are near the maximum
size for the subspecies and would certainly average larger.

I would assume that oressinoma ranges into Ecuador, but I have not seen
Ecuadorian specimens referable to this subspecies. It is replaced to the
south by quitensis Nelson, which differs from oressinoma in having the cap

104 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

more extensive, the cap and underparts more deeply ochraceous, as well as
averaging larger. Hellmayr (1936) considered quitensis to be doubtfully dis-
tinct from brevirostris east of the Andes, but the former is a larger bird with
a markedly longer bill and a less intensely colored cap.

Euphonia xanthogaster brevirostris Bonaparte

Euphonia brevirostris Bonaparte 1851, Rev. Mag. Zool. 2(3):136. **Colum-
bia’’ = Bogota.

Characaters.—Differs from chocoensis and oressinoma in having the un-
derparts and cap darker, more ochraceous, less yellow. Cap plusher, more
velvety, without noticeable dark bases to the feathers, hence mostly un-
spotted. Bill smaller than in oressinoma.

Range.—In Colombia found on the eastern slope of the eastern Andes
from Cundinamarca southward, and on the southern end of the eastern slope
of the central Andes in Huila. The stated range also includes eastern Ec-
uador and Peru, southern Venezuela, the Guianas, and northwestern Brazil.

Specimens examined.—COLOMBIA. CUNDINAMARCA: ‘‘Bogota”’ (6,
USNM); vicinity of Bogota (1, USNM); near Quetame (1, ANMH). META:
Buena Vista (1, ANSP); Mt. Macarena (2, AMNH). CAQUETA: Rio Ca-
queta (1, USNM); La Murelia [=Morelia], Rio Bodoquera (1, USNM; 3,
AMNH; 1, ANSP); Florencia (1, AMNH). PUTUMAYO: Rio Rumiyaco
(2, ANSP); Rio Churo Yacu (3, ANSP). NARINO: Rio San Miguel (5,
ANSP). HUILA: La Candela, 10 mi (16 km) SW of San Agustin (8, USNM;
1, AMNH; 16, ANSP); Belen, 45 km SW of La Plata (3, USNM), La Palma
(1, AMNH); Caqueta Trail (1, ANMH); El Isno (1, ANSP). Specimens from
elsewhere in the range were examined but not studied in detail.

Remarks.—Records of this subspecies from Santander del Norte (Meyer
de Schauensee, 1951) are erroneous, being based on intergrades between
E. x. oressinoma and E. x. badissima (q.v.). Thus there appears to be no
evidence that this subspecies occurs any farther north in the eastern Andes
than the Bogota region.

Euphonia xanthogaster exsul Berlepsch

Euphonia xanthogaster exsul Berlepsch 1912, Verh. V Intern. Ornith.
Kongr. Berlin 1911, p. 1017. San Esteban, near Puerto Cabello, Carabobo,
Venezuela.

Characters.—Markedly distinct from any of the preceding subspecies.
Adult males with cap rich orangish chestnut; breast and midline of belly
very deep fulvous, almost chestnut. Females much buffier on breast and
undertail coverts, cap suffused with chestnut, not yellowish.

Range.—Mountains of northern Venezuela from Lara east to Miranda.

VOLUME 94, NUMBER 1 105

Specimens examined.—VENEZUELA. LARA: mountains near Bucari-
to, Tucuyo (1, AMNH). YARACUY: Lagunita de Aroa (1, CM); Aroa,
Bolivar RR (1, CM). CARABOBO: San Esteban, inland of Puerto Cabello
(2, AMNH); La Cumbre de Valencia (1, CM); La Cumbre (1, CM); Las
Quigas (2, CM). ARAGUA: Rancho Grande (2, USNM). DISTRITO FED-
ERAL: El Limon (1, USNM; 2, CM). MIRANDA: Santa Lucia (1, USNM;
1, CM). 7>MONAGAS: Caripé (1, AMNH) [Hellmayr (1936) questioned the
provenance of this specimen and I have not included the locality in Fig. 1].

Remarks.—Birds from the Sierra Periyja, Santander del Norte, and Bo-
yaca, previously referred to exsul, belong to the following new subspecies.

Euphonia xanthogaster badissima, new subspecies

Holotype.—USNM 375042, adult male, Bellavista, “‘pumping station 100
km from Petrolea’’ [=65 km NW of Petrolea as marked on map of Carriker
localities on file at USNM], Santander del Norte, Colombia. Collected 4
July 1943 by M. A. Carriker, Jr., original number 4595.

Characters.—Like exsul but adult males with chestnut of cap darker, less
orangish, fulvous portions of underparts darker and somewhat more exten-
sive. In females the cap is darker and more restricted than in exsul. Males
are very similar to the widely disjunct ruficeps Lafresnaye and D’Orbigny
1834 of Bolivia; they do not differ in the color of the cap but are more
extensively and deeper fulvous below.

Range.—Colombia in the Sierra Perija area of Magdalena, south to San-
tander del Norte, including the Tachira Valley, and on the east slope of the
eastern Andes in Boyaca.

Specimens examined.-COLOMBIA. MAGDALENA: Monte Elias, Sier-
ra Negra, SE of Fonseca (1, USNM); Tierra Nueva, Sierra Negra, SE of
Fonseca (1, USNM). SANTANDER DEL NORTE: Palo Gordo, 10 mi (16
km) SE of Villa Felisa, Tachira Valley (1, USNM); Bellavista, 65 km NW
of Petrolea (3, USNM). BOYACA: Rio Negro (2, CM).

Etymology.— ‘Most chestnut,’’ from Latin badius, chestnut brown, and
-issimus, an adjectival superlative.

Remarks.—A male and female from Rio Negro, Boyaca, are clearly more
similar to badissma than to exsul, although the underparts of the male are
lighter than in the other specimens of badissima.

Euphonia xanthogaster oressinoma X Euphonia xanthogaster badissima

Characters.—Underparts and cap of adult males more ochraceous than
in oressinoma but not nearly as dark as in badissima. Superficially some-
what similar to brevirostris, but cap not plush and velvety, with bases of
feathers darker, giving a spotted appearance; bill larger.

106 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Range.—Western slope of the eastern Andes of Colombia in Magdalena
and Santander del Norte, west of Ocana.

Specimens examined.—COLOMBIA. MAGDALENA: La Palmita (1,
ANSP; 4, CM; 2, USNM). SANTANDER DEL NORTE: Ocana, beyond
Pueblo Nuevo (1, USNM); Pueblo Nuevo (1, ANSP).

Remarks.—These birds had previously been identified as brevirostris
(Meyer de Schauensee, 1951), but they are clearly intergrades between the
two disparate subspecies listed above, whereas the nearest known occur-
rence of brevirostris is from far to the south, on the other side of the eastern
Andes. All of the intergrades come from a rather limited area west of Ocana.
The male labelled ‘““beyond Pueblo Nuevo’’ comes from slightly farther to
the east than those from Palmita and has a darker crown, hence tending
more towards badissima. The specimen labelled simply **“Pueblo Nuevo,”’
however, looks like those from Palmita.

Acknowledgments

This revision is based on specimens in the collections of the following
institutions: American Museum of Natural History, New York (AMNH);
Academy of Natural Sciences of Philadelphia (ANSP); Carnegie Museum
of Natural History, Pittsburgh (CM); National Museum of Natural History,
Smithsonian Institution, Washington (USNM). I am grateful to Frank B.
Gill, Ivy Kuspit, and Kenneth C. Parkes for facilitating this study. This is
contribution number 7 of the Wetmore Papers, a project supported in part
by trust funds from the Smithsonian Institution for completing unfinished
work and study of undescribed material left by the late Alexander Wetmore.

Literature Cited

Hellmayr, C. E. 1936. Catalogue of Birds of the Americas. Part 9.—Field Museum of Natural
History, Zoological Series 13(9):1—458.

Meyer de Schauensee, R. 1951. The birds of the Republic of Colombia. Part 4.—Caldasia
Ses/ =U.

National Museum of Natural History, Smithsonian Institution, Washing-
ton, D.C. 20560.

Note added in proof: The record of Euphonia xanthogaster from El
Real, Darien, Panama, reported fide Wetmore in Haffer (1975, Bonner
Zoologische Monographien 7:158—9), was based on a specimen that I have
re-identified as E. luteicapilla (USNM 484881); this happens to be the
easternmost record of the species.

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 107-121

SPADELLA LEGAZPICHESSI, A NEW BENTHIC
CHAETOGNATH FROM ENEWETAK,
MARSHALL ISLANDS

Angeles Alvarino

Abstract.—Spadella legazpichessi is described from adult mature individ-
uals and compared with related species, S. schizoptera, S. moretonensis,
S. johnstoni, S. sheardi, S. nana, S. pulchella, S. hummelincki, with which
it agrees in the presence of adhesive digital organs. The diagnostic charac-
teristics of these species and S. legazpichessi are compiled in a table to-
gether with information on their geographical distribution.

Spadella is a genus of Chaetognatha including animals of benthic regimen.
These animals are small, from less than | mm to less than | cm. Their
restricted habitat and movements limit the distribution of the species. Spec-
imens of Spadella do not appear in the usual plankton collections, and
collectors of benthic animals usually overlook these small organisms. There-
fore, they are obtained only occasionally. Up to now the number of species
is one dozen, but I am sure that with proper sampling along the coastal
regions of the world, many new species will be found, and a more adequate
distributional range of all the known species of Spadella will be obtained.

Alvarino (1970) reviewed the geographic distribution of the species. Since
then two more species of Spadella have been described, Spadella brad-
shawi Bieri, 1974 and Spadella gaetanoi Alvarino 1978, inhabiting California
and the Hawaiian Islands, respectively. Also, S. angulata was observed at
the Laccadive Islands (Nair and Rao, 1973). Now specimens of a new Spa-
della have been collected at the lagoon of Enewetak (formerly Eniwetok),
Marshall Islands.

The new species of Spadella described here is related to S. schizoptera,
S. moretonensis, S. johnstoni, S. sheardi, S. nana, S. pulchella, S. hum-
melincki, in the presence of adhesive digital organs on the ventrolateral
sides of the tail segment. However, S. schizoptera, S. johnstoni, and S.
sheardi have been described with two pairs of lateral fins, while S. more-
tonensis, S. nana, S. pulchella, and S. hummelincki have only one pair of
lateral fins. Intestinal diverticula are absent in these species and in Spadella
n. sp., although Yosii and Tokioka (1939) indicated they were present in S.
schizoptera, where they are absent according to Conant (1895), Ritter-Za-
hony (1911), Mawson (1944). Probably the specimens analyzed by Yosii and
Tokioka (1939) do not belong to S. schizoptera.

108 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Spadella legazpichessi new species
Figs. 1, 2

Material.—Holotype (USNM No. 60191), paratypes (USNM No. 60192)
collected at Enewetak atoll lagoon, Marshall Islands, in May 1979.

Diagnosis.—The anatomical features described here are based on mature
adult specimens.

The body is opaque, with well developed muscles on the dorsal and ven-
tral sides. Lateral sides very narrow. The whole animal presents a peculiar
iridescence characteristic of most Chaetognatha. No pigment or pigmenta-
tion patterns have been observed (Fig. 1A, B).

Total length when mature, up to 2 mm, tail fin not included. The body is
widest at the region of the transverse septum separating trunk and tail seg-
ments.

The head is large, strong, about the same length and width, but wider
than the neck. It is about 13% of the total length of the animal. The neck
is distinct and covered by a thick collarette (Fig. 2A).

The caudal segment constitutes 50% of the total length of the animal.

The eyes are large, oval, placed at center of dorsal side of the head, and
at about the same distance from each other or slightly closer to each other
than to the sides of the head. The pigmented region is large, formed by 2
branches; the longer extending parallel to the longitudinal axis of the body
and curving laterally at each end; it is crossed at midlength by a short, thick,
bifurcated transverse branch that runs medially. The pattern of the pigment
in the eyes of Spadella legazpichessi is similar to that found in most species
of Spadella. The pigment separates 3 large and one small clear amber
spaces, filled by lenses. The bifurcation of the transverse branch of pigment
is at times quite deep (Fig. 2A).

The hooks are strong, thick, strongly curved, as in Sagitta ferox and S.
robusta, usually a group of 8 or 9 at each side of the head. The anterior
teeth are 3 to 5 per set, long curved inward and posteriorly. The second and
third teeth at each side, counting from the center of the head, are longer
than the others, the third being the longest; the fifth is the shortest. No
posterior sets of teeth were observed.

The corona ciliata is ring-shaped, with the transverse axis slightly longer
than the longitudinal axis. It is located at the dorsal side of the neck, ex-
tending into the collarette region of the neck (Figs. 1A and 2A).

The collarette extends in a thick stratum from the head to level of pos-
terior septum. It is thickest at the neck, tapering towards the posterior part
of the trunk, and extending as a narrow stratum to the tip of the tail.

Intestinal diverticula absent.

The ventral ganglion is large, thick, occupying totally the width of the

109

VOLUME 94, NUMBER 1

0.1mm

Spadella legazpichessi: A, Dorsal view; B, Ventral view.

Fig. 1.

110

0.1mm

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

¢ Ma SW)

0.1mm

Fig. 2. Spadella legazpichessi: A, Dorsal view of head, with detail of hooks, teeth, eyes,
corona ciliata, neck, collarette, and anterior part of intestine; B, Dorsal view of posterior part

VOLUME 94, NUMBER 1 111

trunk and half the length of the trunk segment. It is located at midlength of
the trunk, as close to the neck as to the posterior septum (Fig. 1B).

There is one pair of lateral fins, extending from a level on the trunk
anterior to the opening of the oviducts, to the seminal vesicles. They are
completely rayed. The caudal fin is long, spatula-shaped, rounded laterally
along the tip of the tail segment, with a straight posterior edge. It is not
continuous with the lateral fins, and starts at posterior edge of the seminal
vesicles. The portion of the tail segment surrounded by the tail fin is about
16% of the total length of the animal (Fig. 1A, B).

Adhesive digital organs are formed by three long rigid finger-like pro-
cesses covered by a few small thin papillae. They extend along the ventro-
lateral side of the tail segment. The most anterior finger-like structure has
part of the anterior edge joined to the posterior edge of the lateral fins, but
the other 2 fingers are free. The posterior finger is the longest, and the
anterior one is the shortest. The adhesive organs are at the ventrolateral
side of the tail segment, exactly at the level of the seminal vesicles. Thus
in the tail segment the seminal vesicles are dorsolateral, while the adhesive
organs are ventrolateral (Fig. 2C, D). The adhesive organs are strengthened
by long thin rays or muscle-like fibers, similar to the fin rays. The tip of the
longest finger when extended, reached half-way between the seminal vesi-
cles and the tip of the tail segment. These finger-like organs extend ventrally
to support the animal away from the substratum while crawling, or to help
in fastening the animal strongly to the substratum during the crawling mo-
tion, while the fins are used during the darting and swimming activities. In
the dozens of specimens analyzed no variability was observed in the ad-
hesive digital organs, such as that found by Feigenbaum (1976) in S. schi-
zoptera reared in the laboratory. Such variability might be an artifact of the
culture environment, not occurring in nature. However, adhesive organs in
young specimens are not fully developed, the anterior finger appears first
and the other two at each side of the animal are already incipient.

A cup-like structure is located ventrally at each side of the tail between
the origin of the tail fin and the seminal vesicles. These cups may have
additional adhesive function from the papillae covering the cup process and
the secreted mucus (Fig. 2C, D).

The ovaries extend to the neck region. The ova are few and large, and
they press laterally against the intestine, which usually runs in an S-shape

of tail segment, with detail of seminal vesicles and tail fin. The left seminal vesicle is bursting;
C, Ventral view of posterior part of tail segment, with detail of seminal vesicles, adhesive
digital organs, and tail fin; D, Left side view of posterior part of tail segment, showing the
dorsolateral position of seminal vesicles, and ventrolateral position of adhesive digital organs.

112

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Differential characteristics of mature adult species of Spadella with digital adhe-

sive organs.

Species

Charact.

Body length
(mm)

Head

Tail segment
% to total
length

Lateral
paired fins

S. schizoptera
Conant 1895

1.9-4.9; wider
at level of
posterior
septum

Broader than
body with
narrow neck.
A pair of
pads at
mouth.

47.0—53.7

2 pairs; anterior
pair
extending
from a point
slightly
behind the
ventral
ganglion to
opening of
oviducts. The
second pair
from this
point to
seminal
vesicles.

1 pair, from

S. moretonensis

Johnston and S. johnstoni

Taylor 1919 Mawson 1944
3.68 4.6; slender,
yellow with
brownish
spots on

corona ciliata

Broader than

long. Neck
conspicuous.

56.5; two club- $2.0

shape
papillate
bodies on
posterior part
of ventral
side

Similar to S.
oviducts to schizoptera
tail fin.

Completely

rayed and

covered with

sensory

spots.

S. sheardi
Mawson 1944

3.9-6.5; opaque,
mauve
brownish
pigment along 3
longitudinal
and 2
transverse
bands. At level
of oviducts,
yellow spots on
body.

44.8

2 pairs; anterior
short, on trunk.
Posterior from
opening of
oviducts to
seminal
vesicles.

VOLUME 94, NUMBER | 113

Table 1.—Continued.

S. nana S. pulchella S. hummelincki S. legazpichessi
Owre 1963 Owre 1963 Alvarino 1970 Alvarino n. sp.
0.75—2.40 1.9-2.7; clusters of 2.0—3.0; broadest at 1.8—2.2; thick,
redish cells at the trunk strong, Opaque,
anal region widest at mid-
length.
Broader than Slightly broader Large, roundish; Large, roundish.
widest part of than widest part smaller than in Neck well
body. Neck of trunk. Neck S. pulchella. distinct.
distinct. A pair of distinct. A pair of Neck thick but
papillae between prominences as distinct.
anterior teeth and in S. nana but
mouth with fewer
papillae.
40-50 $2-—55 49 50
1 pair; from a level 1 pair; from a point 1 pair, long, | pair, extending
anterior to anterior but close narrow, from from posterior
opening of to oviducts to level anterior to part of trunk to
oviducts to seminal vesicles opening of seminal vesicles.
seminal vesicles. extending oviducts to
ventrally over seminal vesicles
seminal vesicles extending
joining tail fin ventrally over

seminal vesicles
joining tail fin

114 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

S. moretonensis

Species S. schizoptera Johnston and S. johnstoni S. sheardi
Charact. Conant 1895 Taylor 1919 Mawson 1944 Mawson 1944
Caudal fin Long, spatula Spatula shape, Spatula shape
shape completely
rayed,
covered with
sensory
spots.
Eyes Large, with no Small, widely
pigment spaced and
overlain by
brown pigment
Hooks 8-11, long, 9, slightly 10-11 Up to 11
curved. curved like in
Points sharp E. hamata
Anterior 2-3 long, 3-4, stout, 2, long, about 3, long, about 4
teeth slender curved. half length of to 1% length of
curved hooks. hooks.
towards the
midline where
they meet.
Posterior None None None None
teeth
Corona Three corned Roughly Elliptical- Triangular
ciliata shape or elliptical, rectangular,
pear-shaped slightly mainly on
to roughly pointed at neck.
triangular. anterior end.
Part on head
and part on
neck.
Ventral Large, thick, Large at
ganglion overlaid by anterior half
numerous of trunk.
sensory

spots.

VOLUME 94, NUMBER 1

Table 1.—Continued.

S. nana
Owre 1963

S. pulchella
Owre 1963

S. hummelincki
Alvarino 1970

S. legazpichessi
Alvarino n. sp.

Spatula shape,
Starting at
posterior end of
seminal vesicles

5-9

Spatula shape
starting at a
distance of
posterior end of
seminal vesicles.
This distance is
equal to the
length of the
vesicle.

The pigment ina
longitudinal band
crossed by a
short band at the
center.

8-10

Long spatulated
shape, starting at
a distance of
posterior end of
seminal vesicles
equal to half the
length of the
vesicle. No
rayless zone.

Large, roundish.
Larger than in S.
pulchella.
Pigment in a
thick X-shape.

8—9 each side.
Slender, slightly
curved.

Spatulate in shape,
Starting at
posterior end of
seminal vesicles.
Covered by 12
sensorial spots.
No rayless zone.

Large, oval.
Pigment in a
longitudinal band
tilted towards the
sides, and at
midlength a short
band towards the
center

8-9 at each side.
Strong, curved
like in S. ferox

1-3, the innermost

longer than the
others. Curved

2—4, long, slender,
curved.

4 at each side.
Long, thin, bent
at about

Up to 5 at each
side. First 3
longer than the

towards the midlength. others. The 3rd
midline. at each side is
the longest.
Curved towards
the ventral side.
None None None None

Variable in shape.
Located at the
neck extending
towards the
head.

As broad as long.

Broad, irregular
oval, located at
neck and
extending
towards the
head.

Large, midway
between
posterior edge of
corona ciliata and
posterior septum.

On neck, roundish
with peak
towards center of
head.

Large, thick, about
half length of
trunk, and almost
as wide as trunk.
Closer to
posterior septum
than to neck.

Like a ring at
dorsal part of
neck.

Large, thick at
midlength of
trunk, and about
50% length of
trunk. As close
to neck as to
posterior septum.

116 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Species S. schizoptera

S. moretonensis
Johnston and

S. johnstoni

S. sheardi

Charact. Conant 1895 Taylor 1919 Mawson 1944 Mawson 1944
Sensorial Abundant, On tail fin, Arranged in two
spots extending on lateral fins, symmetric
longitudinal sides of body groups along
and and head. the body.
transverse
rows.

Ovaries Reaching neck Reaching to Up to neck. Open
or anterior level of at clear space
end of ventral ventral between
ganglion or ganglion. Few posterior end
its midlength. large ova. of anterior fin

Opening of and anterior
oviducts end of
swollen and posterior fin.
trilobed.

Seminal Ellipsoidal, Small, Oval, yellow in

vesicles reniform, inconspicuous, color in living
touching both on posterior specimens.
lateral and third of tail
tail fins. segment.
Intestinal Present or Absent
diverticula absent (1)
Adhesive Hand shaped, Rudimentary. At ventral side Laterally at
organs prolongation of seminal ventral side tail
of lateral fins vesicles, between
with 4, 5, 6 extending posterior fins
finger-like over length of and seminal
processes, tail fin. vesicles,
extending arranged in 2
from groups at each

posterior end

of lateral fins.

First finger
longest and
outmost
shortest. All

side, one
towards the
anterior part of
the animal and
other towards
the end. 10-11

VOLUME 94, NUMBER 1 Li 7/

Table 1.—Continued.

S. nana S. pulchella S. hummelincki S. legazpichessi
Owre 1963 Owre 1963 Alvarino 1970 Alvarino n. sp.
Arranged Arranged Numerous Symmetrically

symmetrically as in
S. nana. Lateral

arranged on tail
fin (6 at each

symmetrically in
pairs on caudal

fin and spots on outer side) and 6
longitudinal rows edges of conspicuous
on the body. collarette and along edge of

lateral fins and
more than 6 at
each side along
edges of
collarette.

lateral fins are
conspicuous.

Reach to neck or
anterior end of

Reach neck region.
Few large ova,

Up to midlength of
trunk, 2-3 large

Reaching to neck.
2—5 huge ova and

intestine is Ova. ventral ganglion. pushing intestine
pushed into an S- Few large ova. into an S-shape.
shape.

Oval, roundish Oval, extending Oval or pear Ellipsoidal,
anterior to tail from posterior shaped, touching reniform,
fin. Protected end _ of lateral fins posterior end of touching

posterior end of
lateral fins and
anterior end of
tail fin. Open at
middle of edge
towards posterior
half of vesicle.

Absent

lateral fins to the
prolongations of adhesive organs
lateral and tail and apart from
fins. tail fin. Open

by lateral dorsal
slit at about
midlength.

Absent

to adhesive
organs.

ventrally by

Absent Absent

Extending ventrally Extending from One hand shaped at__— Three long thin

from posterior
end of lateral fins
to seminal
vesicles. Divided
in 2 stout distally
tubercular
fingers, with
muscular fibers.
First finger is
longest. In small
specimens, only
one finger per
organ.

posterior end of
seminal vesicles
to the point the
caudal fin starts.
Not associated
with lateral fins,
not attached to
caudal fin. 2-3
fingers in each
organ. The
innermost finger
longer, slender,
undivided; the

each side, with 3
short, thick stout
fingers each
covered by thin
papillae. Appear
as extension of
lateral fins,
strengthened by
ray-like pattern,
continuation of
the lateral fins
and rising from
latero-ventral

fingers in each
organ,
prolongations of
posterior end of
lateral fins, but
attached also at
latero-ventral
part of tail at
level of seminal
vesicles only.
Not attached to
tail fin.
Outermost finger

118 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

S. moretonensis

Species S. schizoptera Johnston and S. johnstoni S. sheardi
Charact. Conant 1895 Taylor 1919 Mawson 1944 Mawson i944
have adhesive fingers on each
papillae. A side, covered
sensory spot with papillae.
at base of
outermost
branch.
Collarette Thick at neck, Thick at neck
extending reaching
along base of lateral fins.
lateral fins to Thus neck
seminal appears wider
vesicles. than the
head.
Geographic Bahamas, Queensland New South New South
distribution Florida, New Wales Wales.
South Wales?
Misaki
(Japan)?

(1) Characteristic varies with authors (see Alvarino, 1970).

from neck to anus. The opening of the oviducts are dorsolateral cups with
an outer rim. The lateral fins extend ventrally anterior to the openings of
the oviducts. The ventral disposition of the fins and dorsal placement of
oviducts constitute a protective functional device to help in the transference
of sperm during the copulatory process.

The seminal vesicles touch both posterior end of lateral fins and anterior
end of caudal fin. They are large, reniform, about 8% of the total length of
the animal. They open at the dorsal side by a dorsolateral slit in the posterior
half, just beyond the midlength of the vesicle. This opening protrudes to-
ward the posterior part in a short bottle neck, which adapts to the opening
of the oviducts (Fig. 2B, D).

The sensory spots appear distributed on the tail fin, and along the edges

VOLUME 94, NUMBER I

Table 1.—Continued.

S. nana
Owre 1963

Thick at neck,
tapering
progressively
towards the

seminal vesicles.

S. pulchella
Owre 1963

other may split at
the tip. Abundant
papillae cover tip
of2 Tons
digitations.

Very thick at neck

and anterior part
of trunk,
extending along
base of lateral

S. hummelincki
Alvarino 1970

side of tail
segment between
posterior end of
seminal vesicles
and start of tail
fin. Attached by
a point at tail fin,
at the sensory
club.

Wide, thick,

extending from
head to posterior
septum, widest at
anterior half of

119

S. legazpichessi

Alvarino n. sp.
is the longest,
reaching
midlength from
vesicle to tip of
tail. Fingers
strengthened by
thin muscular
fibers. Large
sensory club on
ventral side of
tail at posterior
end of seminal
vesicles.

Thick at neck

extending along
lateral part of
trunk, tapering
towards posterior

At head appears fins to posterior trunk. part of tail
continuous with septum. segment.
hood.
Florida Puerto Rico Southwest of Bahia Enewetak (Marshall
Fosforescente, Islands)
Puerto Rico

of the lateral fins and edges of the collarette, from the head to the posterior
part of the trunk. In the tail fin are 6 at each side, 6 along the edge of each
lateral fin, and more than 6 along the edge of the collarette, from the point
the collarette joins the anterior edge of the lateral fins to the head.
Remarks.—Spadella legazpichessi differs from S$. schizoptera, S. john-
stoni, S. sheardi in many anatomical characters, especially in the number
of lateral fins, from S. schizoptera probably also in the intestinal diverticula,
and from S. moretonensis, S. nana, S. pulchella, and S. hummelincki main-
ly in the shape of hooks, head, characteristics of the adhesive organs, po-
sition of ventral ganglion, and position of seminal vesicles (Table 1).
Distribution.—The specimens studied were collected at Enewetak atoll,
Marshall Islands, on sand, rubble coral, and living coral. The collections

120 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

were obtained with a bottom emergence trap for epibenthic and hypopelagic
organisms, during studies on benthic marine communities. In those collec-
tions were also trapped pelagic species of Chaetognatha, Sagitta ferox, S.
robusta, S. neglecta, S. oceania, which might be swimming and feeding in
the water stratum over the bottom.

Etymology.—Named after the navigator Miguel Lopez de Legazpi, whose
party discovered the Marshall Islands in 1565, then named “‘Islands of
Bearded people,’ and James R. (Tony) Chess, who kindly provided me
with the material for this study. Legazpi’s expedition reached those islands
and atolls with four sailing vessels under Legazpi’s Command (San Pedro,
San Pablo, San Juan and San Lucas) departing from Navidad port (north
of Acapulco) on November 1564 and reaching those islands, coral reefs, and
atolls in January 1565, in their route to the Philippines. Legazpi’s expedition
constitutes the initiation of the first round trip across the Pacific, inaugu-
rating with their return from the Philippines to Acapulco, the first regular
transoceanic route, which also lasted longest in history, 1564 to 1815 (Co-
leccion de Diarios y Relaciones para la Historia de los Viajes y Descubri-
mientos).

Acknowledgments

I wish to express my gratitude to Dr. Thomas E. Bowman for his excellent
editorial suggestions to improve the clarity of the work, and also my appre-
ciation to James R. Chess for sending me the material for this study and
Dr. John R. Hunter for reading the manuscript and for valuable comments.

Literature Cited

Alvarino, A. 1970. A new species of Spadella (Benthic Chaetognatha).—Studies on the Fauna

of Curacao and other Caribbean Islands 24:73-89.

. 1978. Spadella gaetanoi, a new benthic Chaetognath from Hawaii.—Proc. Biol. Soc.

Washington 91(3):650-657.

Bieri, R. 1974. A new species of Spadella (Chaetognatha) from California.—Publ. Seto Mar.
Biol. Lab. 21(3—4):281-286.

Conant, F. S. 1895. Description of two new Chaetognaths (Spadella schizoptera and Sagitta
hispida).—Johns Hopkins Univ. Circ. 14(119):77-78.

Feigenbaum, D. L. 1976. Development of the adhesive organ in Spadella schizoptera (Chae-
tognatha) with comments on growth and pigmentation.—Bull. Mar. Sci. 26(4):600—603.

Johnston, T. H., and B. B. Taylor. 1919. Notes on Australian Chaetognaths.—Proc. Roy.
Soc. Queensland 31(3):28—41.

Mawson, P. M. 1944. Some species of the Chaetognatha genus Spadella from New South
Wales.—Trans. Roy. Soc. South Australia 68(2):327-333.

Nair, V. R., and T. S. S. Rao. 1973. Chaetognaths from the Laccadives with the new record
of Spadella angulata (Tokioka, 1951). Pp. 319-327 in: Bernt Zeitschel, ed., The Biology
of the Indian Ocean, XIII + 549 pp., Springer-Verlag, New York, Heidelberg, Berlin.

Owre, H. B. 1963. The genus Spadella (Chaetognatha) in the western North Atlantic Ocean,

VOLUME 94, NUMBER | 121
with descriptions of two new species.—Bull. Mar. Sci. Gulf and Caribbean 13(3):378-

390.
Ritter-Zahony, R. 1911. Revision der Chatognathen. Deutsch.Stidpolar-Exped. 1901-1903, 13,

Zool. 5(1):1-71.
Yosii, N., and T. Tokioka. 1939. Notes on Japanese Spadella (Chaetognatha).—Annot. Zool.

Japan 18(4):267-275.

National Marine Fisheries Service, NOAA, Southwest Fisheries Center,
P.O. Box 271, La Jolla, California 92038.

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 122-134

GORGONISCUS INCISODACTYLUS, A NEW ISOPOD
OF THE EPICARIDEAN SUPERFAMILY
CRYPTONISCOIDEA, PARASITIC ON AN

ASCOTHORACICAN CIRRIPED FROM HAWAITI

Mark J. Grygier

Abstract.—A description of a cryptoniscoid isopod, Gorgoniscus inci-
sodactylus, n. gen. et sp., is presented. This is the first isopod parasite to
be described from an ascothoracican cirriped and the first cryptoniscoid
from Hawaii. The new species is compared to representatives of other cir-
riped-infesting cryptoniscoid families, but its familial status remains uncer-
tain. The significance of the first antennae, mandibles, pleopodal chitinous
rings, and uropod setation of the cryptoniscus larva, and the posterior struc-
tures, and cuticular spicules of the female is discussed. It is suggested that
this isopod is a parasitic castrator or an egg parasite, and that observed high
infestation rates (up to 67%) may affect the reproductive success of the host
population.

Cryptoniscoidea is a superfamily of epicaridean isopods, various members
of which are parasites of peracarids, ostracods, cirripeds (thoracicans and
rhizocephalans), and copepods (as larvae). They are protandric hermaph-
rodites, and have a complex life cycle usually involving 2 larval stages
parasitic on copepods, the ““cryptoniscus’’ larva which seeks out the defin-
itive host, the protandric male indistinguishable from the cryptoniscus larva,
and the female, derived by more or less catastrophic metamorphosis from
the protander. The term “‘cryptoniscus larva’’ will be used herein to refer
to either the larva or the protander, while the term ‘‘cryptoniscoid’’ will
refer to the superfamily as a whole. The cryptoniscus larva is relatively
unmodified; the female, however, is commonly simplified to a large brood
sac, though sometimes part of the cryptoniscus larval externum is retained.

Two published reports exist of isopods parasitic upon ascothoracican cir-
ripeds (Ascothoracica is a primitive group of crustaceans whose members
are parasites of echinoderms and anthozoans). Newman (1974) mentioned
in his description of Synagoga sandersi that a gravid female was host to an
isopod occupying the space of an egg. The isopod was given to Newman’s
student, the late Larry Ritchie, for study; it has not yet been possible to
locate the specimen, but Newman is certain that it was a cryptoniscoid
(personal communication). Pyefinch (1939) reported the discovery of cryp-
toniscoid parasites from three species of the zoanthid-infesting ascotho-

VOLUME 94, NUMBER 1 123

racican Baccalaureus. These specimens have not yet been described, but
they are to be included in the Isopoda volume of the John Murray Expe-
dition Reports (Boxshall, personal communication).

Gorgonians (USNM Acc. no. 330459) were collected by the Star IT sub-
mersible at a site off Makapuu Point, Oahu, Hawaii, at 366 meters depth on
27 January 1978. Galls from specimens of several paramuriceid genera pre-
sumably containing barnacles were sent to Scripps Institution of Oceanog-
raphy for identification. These galls contained representatives of the cirriped
order Ascothoracica, which have been identified as a new species of Gor-
gonolaureus Utinomi, 1962. A description of the ascothoracican is in prep-
aration. Several apparently mature, but not gravid specimens of Gorgono-
laureus contained cryptoniscoid isopods in the dorsal brood chamber
formed by the fusion of the carapace valves. A Placogorgia with Gorgon-
olaureus galls was collected at a later date at the same locality, and was
dried. Some of the ascothoracicans from this gorgonian were also infested
with cryptoniscoid isopods, including 2 post-metamorphic females.

The families of Cryptoniscoidea have historically been delimited by their
hosts. However, a revision of these isopods is in progress (Stromberg, per-
sonal communication), so I will not now consider a new family for the
present species despite the novel taxonomic position of its host. Nonethe-
less, it will be compared to representatives of those cryptoniscoid families
known to parasitize cirripeds: Liriopsidae on rhizocephalans and Hemion-
iscidae and Crinoniscinae (formerly Crinoniscidae, but incorporated into
Liriopsidae by Bocquet-Védrine, 1974) on thoracicans.

Isopods were prepared for study as follows. Dried material was recon-
stituted overnight in a 10% trisodium phosphate solution. Most specimens
were studied whole in lactic acid mounts, but | cryptoniscus larva and 1
female were dissected and the parts mounted in Turtox CMC-10 with acid
fuchsin. Drawings were done by the author with the aid of a camera lucida.

Order Isopoda Latreille, 1817
Suborder Epicaridea Latreille, 1831
Superfamily Cryptoniscoidea
(nom. transl. herein pro Cryptoniscinae Bonnier, 1900)
Family incertae sedis
Genus Gorgoniscus, new genus

Diagnosis.—Cryptoniscoid isopod parasitic on ascothoracican cirriped
Gorgonolaureus. Cryptoniscus larva about 0.8—1.0 mm long, fusiform. Eyes
absent. Prominent transverse cuticular striations on dorsum of body. Basal
segment of first antenna with 5 posterior teeth, lateralmost tooth with pro-
nounced shoulder. Second antenna extending posteriorly to third free tho-
racic segment. Oral cone pointing anteriorly; mandibles styliform. Epimeral

124 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

plates of pereon longitudinally ridged, posterior margins entire. Dactyli of
pereopods 6 and 7 longer and narrower than others, claws notched. Pleo-
pods with typical setation, endopodites with 5 setae (3 on fifth pair), ex-
opodites with 5, lateralmost one-third length of others; endopodites with
chitinous ring. Uropod endopodites twice as long as exopodites; endopodite
dorsal setae not necessarily numerically symmetrical. Telson margin entire.

Females about 4 mm long, ellipsoidal to round in dorsal view,:dorsoven-
trally compressed with anterior and posterior ends curled ventrally. Most
appendages absent. Mouthparts varying, either spatulate mandibles or me-
dian proboscis.

Etymology.—From a shortened form of Gorgonolaureus, the ascotho-
racican host of the present species and Latin oniscus (= wood louse). Gender
masculine.

Type-species.—Gorgoniscus incisodactylus, new species.

Gorgoniscus incisodactylus, new species
Figs. 1-4

Material.—Holotype, cryptoniscus larva at U.S. National Museum of
Natural History (USNM 181126); Paratypes: 2 cryptoniscus larvae (1 dis-
sected and mounted, USNM 181127-8); 2 cryptoniscus larvae at Museum
of Comparative Zoology, Harvard University; | cryptoniscus larva retained
at Scripps Institution of Oceanography; 2 2° (1 dissected and mounted,
USNM 181129-30).

Type-locality.—Off Makapuu Point, Oahu, Hawai (21°18’N, 157°32’W)
at 366 meters depth, collected by B. Bartko and K. Muzik in Star IT sub-
mersible 27 January 1978.

Distribution.—Type-locality and nearby location 10 kilometers off Mak-
apuu Point, where B. Bartko collected more specimens at 366 meters depth
with Star IT submersible in 1979.

Host.—Found within brood chamber formed by carapace of Gorgono-
laureus sp. (Cirripedia: Ascothoracica) living in galls on paramuriceid gor-
gonians. See Table | for distribution of isopods on Gorgonolaureus and
various gorgonians.

Etymology.—From Latin inciso (=to cut in) and Latin dactylus (=digit),
referring to the notched claws of pereopods 6 and 7.

Description.—Characteristics of genus with following amplifications.

Cryptoniscus larva (Figs. 1-3): Dorsal profile (Fig. 1A) tapering narrowly
posteriorly, more broadly anteriorly, cephalon bluntly pointed. Length (not
including uropods) 0.84—1.00 mm (Holotype longest); maximum width (at
midlength) 0.25—0.35 mm (Holotype widest). Cross section of body convex
dorsally, flat ventrally, with slight ventral curvature of body longitudinally.
Body (Fig. 1A, B) divided into cephalon with 2 pairs of antennae and oral

VOLUME 94, NUMBER 1 125

Table 1.—Distribution of Gorgoniscus incisodactylus on its host, the ascothoracican Gor-
gonolaureus sp., from various gorgonians. The Placogorgia was collected in 1979, the other
three gorgonians in January 1978.

Gorgonian Gorgonolaureus Isopods
Paramuricea or 1 mature 2 cryptoniscus larvae
Placogorgia (including Holotype)
1 immature none
1 probably immature not examined
Villogorgia 1 mature none
1 mature none
1 nearly mature none
Muriceides 1 mature none
1 mature none
1 mature none
Placogorgia 1 nearly mature 1 cryptoniscus larva
1 mature 3 cryptoniscus larvae
1 mature 12 (first female)
1 mature 12 (second female)
1 cryptoniscus larva (lost)
1 immature none
1 mature none

cone, 7 free thoracic segments, seventh longer than others, each bearing
pair of pereopods, 6 abdominal segments with 5 pairs of pleopods and | pair
of uropods, and telson. Anterior end of body dorso-ventrally striated (Fig.
1C). Color white to gray in preserved specimens.

First antenna (Fig. 1D) not extending past cephalon; peduncle triarticu-
late, distal segment bearing 2 rami. Basal segment large (0.11 mm long);
medial spines absent; medial posterior teeth originating more proximally
than lateral teeth; teeth simple except for lateralmost; this tooth with 1 short
seta at base, | at shoulder, | just proximal to shoulder; ventral surface of
first segment covered with cuticular plates (Fig. 1C); 3 setae near anterior
end, longest one medial, shorter 2 of subequal length; single large spine on
dorsal surface between bases of all teeth. Second segment cylindrical, small-
er than ‘‘palm’”’ of first, bearing 3 setae on lateral side ventrally, 2 long, i
short; possibly another long seta on medial side; large spine on dorsal side
overlapping lateral edge of first segment. Third segment very short, arising
from dorsal side of second; bundle of 14 aesthetascs apparently arising from
it; also 2 setae, 1 lateral, other medial. Terminal rami uniarticulate; dorsal
one with 4 long terminal setae, 2 aesthetascs apparently arising from shoul-
der near base; ventral one with 2 terminal setae and single terminal spinule.

Second antenna originating beneath teeth of first antenna’s basal segment;

126 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

0.02mm j————=4 D E ¢
0.02 mm 4 F
0.1m ——_==—=| ¢
0.5 0 el AB

Fig. |. Cryptoniscus larva of Gorgoniscus incisodactylus. A, Dorsal view of Holotype,
cephalon and first free thoracic segment showing cuticular striation, uropod setae not shown;
B, Ventral view of Holotype, bases only of pereopods and sympodites only of pleopods shown;
C, Ventral view of cephalon showing positions of first and second antennae; D, Dorsal view
of first antenna, medial posterior tooth missing, setae shown as solid, aesthetascs open; E,
Second antenna, basal segment broken off proximally; F, Oral cone showing mandibles and
posterior oval pieces (remnants of second maxillae); G, Dorsal view of uropods, hatched line
near base representing posterior margin of telson; H, Dorsal view of abdominal segments 5
and 6, showing uropods without setae and telson (stippled).

VOLUME 94, NUMBER I 127

Fig. 2. Appendages of Gorgoniscus incisodactylus cryptoniscus larva. A, Pereopods 3
(above) and 4, showing relation to ridged epimeral plates; B, Detail of distal region of pereopod
5; C, Pereopod 1; D, Pereopod 2; E, Detail of pereopod 2 distal region; F—J, First through fifth
pleopods showing size gradation and endopodite ring, terminal setae not shown; K, Typical
pleopod showing terminal setation, not all setules shown.

4-segmented peduncle and 5-segmented flagellum, extending to third free
thoracic segment (not including terminal setae). Setation apprently not con-
sistent; description based on dissected specimen (Fig. 1E). First and second
segments cylindrical with medial distal extension. No setae on first segment;
1 seta on second. Third segment cylindrical, same length as previous seg-
ments but half as wide, with 3 setae at tip. Fourth segment slightly narrower
than third, bearing 4 setae. Flagellar segments cylindrical, distal ones nar-
rower; first and third segments with 2 setae each; second and fourth with
1 each; 4 terminal setae on fifth segment, 3 quite long; lateral terminal spine
on each flagellar segment, thinner and longer on distal segments.

Oral cone between bases of second antennae. Mouthparts (Fig. IF) en-
closed anteriorly by labrum, more or less exposed posteriorly near tip of
cone. Mandibles arising at sides of cone’s base, extending ventrally and
bending slightly posteriorly; tip roughened with small knobs. Pair of oval
plates at posterior base of cone probably representing second maxillae (Gou-
deau, 1969, 1977).

First 2 pairs of pereopods similar (Fig. 2C—E), with dactylus and propodus
forming subchela. Basis cylindrical, twice as long (0.09 mm) as wide. Is-

128 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

chium two-thirds as long as basis, much narrower, bowed posteriorly. Me-
rus triangular with large seta on lateral corner, small seta distally on medial
side. Carpus an elongate triangle with stout spine distally. Propodus as long
as ischium, attached firmly to long lateral side of carpus, with distal artic-
ulation of dactylus, latter modified as heavy claw. When closed, dactylus
engaging 2 heavy spines on propodus, proximal one bifid, distal one trifid.
Dactylus not opposing terminal spine of carpus, but gap between them
small.

Five other pairs of pereopods similar to one another (Fig. 2A, B). Basis
0.12 mm long, narrow but clavate with expanded distal half. Ischium half
as long as basis with lateral edge convex. Merus triangular with large seta
on latero-distal angle and long, fine seta on medio-distal angle. Carpus also
triangular with stout spine medio-distally. Propodus 0.1 mm long, slender,
distal half bearing 2 stout spines (distal one bifid) opposing dactylus. Dac-
tylus not forming subchela; long and narrow, tapering to slightly recurved
claw at tip; small spine at base of claw more pronounced in posterior pairs.
Pereopods 6 and 7 with dactyli 25% longer and more slender than those of
other pairs (Fig. 3).

Five pairs of pleopods similar to one another, becoming gradually smaller
posteriorly (Fig. 2F—J). Pleopods biramous, with sympodite, exopodite, and
endopodite each uniarticulate. Sympodite triangular, about twice as wide as
long, sharpest angle lateral; 2 setae on medio-distal apex. Endopodite rect-
angular, twice as long as wide, outline slightly convex; 3 short spines on
lateral side; chitinous ring either on anterior surface or within. Exopodite
slightly narrower and longer than endopodite, constricted basally with 3
spinules on medial side. Longer setae of each ramus (Fig. 2K) about 0.15
mm long; setules 0.04 mm long.

Uropods (Fig. 1G) biramous. Sympodite short, disc-shaped; 2 strong setae
at latero-distal corner, medial one 3 times longer than lateral; lateral and
medial sides of sympodite lined with fine hairs. Exopodites 0.04 mm long,
cylindrical with short lateral spine, 2 long setae, and | shorter, narrow seta
distally. Endopodite a tapering cylinder lined medially with fine hairs; few
(3-5?) long and short terminal setae as long but weaker than those of ex-
opodite; several short setae (6 on left, 8 on right of specimen examined for
them) basally on dorsal side. Telson short (Fig. 1H), rear margin sinusoidal,
medial part protruding posteriorly.

First female (Fig. 4A—C): Color in preserved state brownish-yellow. Slight
left-right asymmetry (Fig. 4A, B), 4.55 mm long, 4.22 mm wide. About 10
segments visible dorsally, 3 short ones anteriorly, 7 large ones following.
Bilobed labrum (Fig. 4C) 0.4 mm wide, nearly bisected by posterior incision;
3 translucent regions, | medial and pair lateral. Anteriorly directed probos-
cis 0.25 mm long, tapering distally. Two small papillae, possibly reduced
mandibles or openings of maxillary glands, posterior and lateral to base of

VOLUME 94, NUMBER 1 129

Vix
© < =
TH IV V
=)
0.02mm
| |

Fig. 3. Propodi and dactyli of pereopods 3-7 in cryptoniscus larva of Gorgoniscus inci-
sodactylus, showing longer and narrower dactyli with notched claws in pereopods 6 and 7.

proboscis near posterior margin of labrum. Two featureless flaps of tissue
just posterior to labrum. Other appendages absent. Telson not visible,
though posterior end of body obscured by longitudinal folds in posteriormost
segments (Fig. 4B). Eggs absent.

Second female (Fig. 4D, E): Color in preserved state dull yellow. About
Same size as first female, but more elongate. Anterior end squared off,
hoodlike. Demarcations of few anterior segments visible, but most of body
covered with fuzzy growth. Mouthparts near base of cephalic hood (Fig.
4D), consisting of anterior-pointing labrum and mandibles similar to those
in cryptoniscus larva, but laterally flattened and relatively shorter with nu-
merous small hooks on distal margins. Posterior conical structure (telson?)
with distal depression (anus?) (Fig. 4E); uniarticulate structure at one side

130 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1.0mm

-——A AB

r 0.1 mm SDE
r 0.2 mm i

Fig. 4. Females of Gorgoniscus incisodactylus. A, Dorsal view of first female, anterior end
above; B, Ventral view of first female showing oral region above, longitudinally folded posterior
region below; C, mouthparts of first female showing labrum, proboscis, pair of papillae, and
tissue flaps; D, Mouthparts of second female showing mandibles; E, Posterior body protrusion
of second female with possible rudimentary uropod at base on right.

of base bearing 3 spines posteriorly, | anteriorly. Integument, where not
fuzzy, studded with denticulate spicules in tight groups of 1-4. Eggs absent,
large quantities of lipid or yolk present.

Remarks.—Nielsen and Stromberg (1965, 1973) have reviewed the fami-
lies and genera within Cryptoniscoidea, and, in the latter paper, have listed
pertinent diagnostic features of the cryptoniscus larvae for each family. It
can be shown using these criteria and information from other authors that
Gorgoniscus does not readily fit into any established genus or family of
Cryptoniscoidea. |

Liriopsidae (sensu stricto) have cryptoniscus larvae without dorsal cu-
ticular ridges, with the posterior margin of the first antenna’s basal segment
entire, and with very short propodi on pereopods 6 and 7, conditions con-
trary to those in Gorgoniscus. The mature females in this family have their
body divided into 2 parts connected by a thin waist, and the second per-
eopod is retained well into metamorphosis (Caullery, 1907). This description
does not recall the characteristics of the present females.

Crinoniscinae have cryptoniscus larvae that differ from the present one
only in having an entire posterior margin on the basal segment of the first

VOLUME 94, NUMBER 1 131

antenna. Two genera, Crinoniscus Pérez, 1900, and Leponiscus Gruvel,
1901, have adult females in which the body is a cruciform sac quite unlike
Gorgoniscus (Bocquet-Vedrine and Bocquet, 1972a, 1972b), while a transi-
tional metamorphic form known in Crinoniscus and Proteolepas Darwin,
1854 retains the second pereopods as grasping organs (Bocquet-Védrine,
O72 MOT):

Among Hemioniscidae, only Hemioniscus balani Buchholz, 1866 is well
enough known for comparison. The cryptoniscus larva of this species has
recently been redescribed (Goudeau, 1970). Differences with the Gorgo-
niscus cryptoniscus larva include toothed posterior margins of the thoracic
epimeral plates in H. balani; the latter also has more posterior teeth on the
basal segment of the first antenna. The female retains the anterior portion
(forward of fifth free thoracic segment) of the cryptoniscus larval body but
expands the posterior part into a stellate sac during metamorphosis (Gou-
deau, 1967, 1977), producing a form dissimilar to Gorgoniscus.

The other cryptoniscoid families, Podasconidae, Asconiscidae, Cypron-
iscidae, and Cabiropsidae all parasitize hosts far removed from cirripeds
and have many more differences distinguishing them from the present genus
than have the families discussed above. Finally, according to the compila-
tion of characters given in Nielsen and Stromberg (1973), the combination
of a toothed first antennal basal segment and entire epimeral plate margins
is unique to Gorgoniscus.

It is unlikely that Gorgoniscus can be synonymized with any known genus
of cryptoniscoids. It is less certain whether a new family need be erected
for it. Despite great differences in the morphology of the females, differ-
ences between the present genus and both Crinoniscinae and Hemioniscidae
are minor in the cryptoniscus larval stage. Therefore, it may prove expedient
to incorporate Gorgoniscus into one of these families. Until Stromberg’s
revision is published, it seems wisest to retain the genus separately, incertae
sedis.

Goudeau (1972) showed that the first and second segments of the first
antenna in the cryptoniscus larva of H. balani are independently articulated
to the cephalon, independently movable, and not articulated with each oth-
er. She concluded that the second segment is the actual basal segment. The
first segment (her ‘‘plaque aliforme’’), once part of the first antenna, is now
separated and partly fused to the body, a situation analogous to the trans-
formation of pereopod coxae into epimeral plates. In my dissected speci-
men, the first antenna was removed in one piece. The first segment is evi-
dently articulated to the cephalon near the base of the medial tooth, which
was torn away, but there is no evidence of any articulation of the second
segment directly to the cephalon. Having had no opportunity to observe
live animals, I cannot say whether the segments are independently movable.
If Goudeau’s observations are correct, I agree that the ‘‘plaque aliforme’’

132 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

must be derived from a first antennal segment; perhaps in H. balani a sec-
ondary attachment of the second segment has occurred. Such an arrange-
ment does not seem to be present in G. incisodactylus.

Goudeau (1970, Fig. 9) shows rings in the pleopods of H. balani similar
to those in G. incisodactylus, but she does not mention them in the text.
References to these rings are made in the descriptions of certain other cryp-
toniscoids, but not all. This could be the result of the dearth of complete
descriptions in this group, but Nielsen and Stromberg’s otherwise exem-
plary description of the cryptoniscus larva of a cabiropsid (1965) includes
no mention of these structures. The presence or absence of chitinous rings
in the pleopods may be an unappreciated taxonomic character. I could not
determine whether the rings are external features or internal. If external,
they may be sucking discs for attachment to the host, but setae from anterior
pleopods overlay posterior ones, presumably limiting the rings’ utility for
this purpose. If internal, they may serve as a kind of brace.

The styliform shape and roughened tips of the mandibles suggest that they
are used both for piercing and trituration.

The asymmetry in the number of dorsal setae on the uropod endopodites
is striking, for possession of 6 setae in this position has previously been
considered a conservative feature of the superfamily (Nielsen and Strom-
berg, 1973).

The uniarticulate structure near the base of the posterior protrusion in the
second female may be a rudimentary uropod. There is a scar opposite this
structure which may show the location of the other uropod, ripped off during
dissection. The cuticular spicules are similar to those described by Goudeau
(1974) in H. balani females, though in the latter each spicule is rectangular
in side view with a dentate distal margin. The fuzz covering much of the
second female appears pathological, perhaps a fungal infestation; however,
it is sticky, suggesting a secretion of the isopod, the ascothoracican, or the
gorgonian.

The only Gorgonolaureus specimens infested were mature or nearly ma-
ture females without eggs (Table 1). Maturity was judged on the basis of
comparisons with other, fecund, specimens. This fact suggests that the
isopod is either a parasitic castrator or an egg parasite. The latter possibility
is supported by the fact that isopods were never attached orally to their
hosts, but could be removed with no difficulty from the brood chamber.
There is a large difference in infestation rate between the 2 collection dates.
As Table 1 shows, only 1 of 8 Gorgonolaureus specimens collected in 1978
was infested by G. incisodactylus, while 4 of 6 specimens collected from
Placogorgia in 1979 were parasitized. Though the number of hosts exam-
ined is small, these data suggest that G. incisodactylus may at times have
a large detrimental influence on the reproductive success of its host popu-
lation.

VOLUME 94, NUMBER 1 133

The depth of collection (366 m) is greater than usual for cryptoniscoids,
but other such instances have been reported; e.g., an undescribed deep-sea
ostracod parasite (Nielsen and Stromberg, 1973) and the parasite from Syn-
agoga sandersi, which was collected at 5000 m (Newman, 1974). Finally,
G. incisodactylus is the first cryptoniscoid isopod recorded from Hawaii.

Acknowledgments

I would like to thank Mss. Katherine Muzik and Catharine Kessler of the
Museum of Comparative Zoology for supplying the host ascothoracicans,
Dr. William Newman and Mr. George Wilson of Scripps Institution of
Oceanography for reviewing earlier drafts of this manuscipt, and Dr. Jarl-
Ove Stromberg of Kristineberg Marine Biological Laboratory for helpful
suggestions on the taxonomic treatment. Dr. Newman provided laboratory
space. This work was done during the tenure of a National Science Foun-
dation Graduate Fellowship and is a Contribution of Scripps Institution of
Oceanography, new series.

Literature Cited

Bocquet-Védrine, J. 1972. Suppression de l’ordre des Apodes (Crustacés Cirripedes) et rat-
tachement de son unique représentant, Proteolepas bivincta, a la famille des Crino-
niscidae (Crustacés, Isopodes, Cryptonisciens).—C. R. Acad. Sci. Paris 275D:2145-
2148.
. 1974. Parenté phylogénétique des Isopodes Cryptonisciens rangés jusqu ici dans les
familles des Liriopsidae et des Crinoniscidae, in Arvy, L., ed. Recherches Biologiques
Contempéeraines (Imp. Wagner, Nancy). [Not seen by author. ]
. 1979. Interpretation actuelle de la description de Proteolepas bivincta Darwin, 1854
(représentant unique de l’ancien ordre des Cirripedes Apodes).—Crustaceana 37: 153—
164.
, and C. Bocquet. 1972a. Réalisation de la forme définitive chez Crinoniscus equitans
Pérez, au cours de | étape femelle du cycle de cet Isopode Cryptoniscien.—C. R. Acad.
Sci. Paris 275D:2009-2011.
, and . 1972b. Sur la validité des genres Leponiscus Gruvel, 1901 et Crinoniscus
Pérez, 1900.—Arch. Zool. Exp. Gén. 113:395—400.
Bonnier, J. 1900. Contribution a l’étude des Epicarides, les Bopyridae.—Trav. Statn. zool.
Wimereux 8:1—475. [Not seen by author. ]
Caullery, M. 1907. Recherches sur les Liriopsidae, Epicarides Cryptonisciens parasites des
Rhizocéphales.—Mitth. Zool. Stat. Neapel 18:583-643.

Goudeau, M. 1967. Transformation morphologique du male en femelle chez l’Isopode Epi-
caride Hemioniscus balani Buchholz.—Cah. Biol. Mar. 8:437—448.

1969. Appareil buccal et mécanisme alimentaire chez l’Isopode Epicaride Hemio-
niscus balani Buchholz.—Arch. Zool. Exp. Gén. 110:437-512.

1970. Nouvelle description d’Hemioniscus balani Buchholz, Isopode Epicaride, au
stade de male cryptoniscien.—Arch. Zool. Exp. Gén. 111:411-446.
. 1972. Mode d’articulation a la capsule céphalique et conformation de |’antennule chez
le male cryptoniscien d’Hemioniscus balani Buchholz, Isopode Epicaride.—C. R. Acad.
Sci. Paris 275D: 1997-1999.

134 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1974. Structures cuticulaires chez Hemioniscus balani Buchholz, Isopode

Epicaride.—C. R. Acad. Sci. Paris 278D:3331-3334.

. 1977. Contribution a la biologie d’un crustacé parasite, Hemioniscus balani Buchholz,
Isopode Epicaride. Nutrition, mues et croissance de la femelle et des embryons.—Cah.
Biol. Mar. 18:201—242.

Newman, W. 1974. Two new deep-sea Cirripedia (Ascothoracica and Acrothoracica) from the
Atlantic.—J. Mar. Biol. Ass. U.K. 54:437—456.

Nielsen, S., and J. Stromberg. 1965. A new parasite of Cirolana borealis Lilljeborg belonging

to the Cryptoniscinae (Crustacea Epicaridea).—Sarsia 18:37-62.

, and . 1973. Morphological characters of taxonomic importance in Cryptoniscina

(Isopoda Epicaridea). A scanning electron microscopic study of cryptoniscus larvae.—

Sarsia 52:75—96.

Pyefinch, K. 1939. Ascothoracica (Crustacea, Cirripedia).—Scient. Rep. John Murray Exped.

5(9):247-262.

Scripps Institution of Oceanography A-008, University of California, San
Diego, La Jolla, California 92093.

Note added in proof: The various gorgonians listed in Table | represent a
single undescribed species of Placogorgia (K. Muzik, pers. comm.). Nier-
strasz and Brender a Brandis (1930, Proc. U.S. Natn. Mus. 77(9):1-—9)
described a supposed cryptoniscoid, Faba glabra, from a shrimp collected
near Waikiki, Oahu, so G. incisodactylus may not be the first of this
superfamily from Hawaii, as stated above.

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 135-162

THREE NEW SHRIMPS, AND SOME INTERESTING
NEW RECORDS OF DECAPOD CRUSTACEA FROM A
DEEP-WATER CORAL REEF IN THE
FLORIDA KEYS

Robert H. Gore

Abstract.—A small collection of 55 specimens of decapod crustaceans,
obtained using a research submersible on a deep-water coral reef off Key
Largo, Florida, produced one new genus and three new species of natantian
shrimp, plus several major range extensions to the continental United States
for other decapods. The new taxa included Veleroniopsis kimallynae new
genus and species, and Pontoniopsis paulae, new species, in the Palae-
monidae, and Odontozona libertae, new species, in the Stenopodidae. The
discovery of new taxa in depths of less than 100 m suggests that the cryptic,
coral-associated decapod crustacean fauna may be richer than previously
suspected.

The decapod crustacean fauna of the Florida Keys has long been recog-
nised as being both speciose and numerous. Although no comprehensive
specific inventory has ever been conducted, the fauna is considered rela-
tively well known as a result of several regional and monographic studies
(e.g. Rathbun’s monographs on brachyurans), which included species whose
ranges extended either to, or north- or southward through the Florida Keys.
As might be expected, the intertidal and shallow subtidal decapods are rel-
atively better known than their deeper-living counterparts, because of the
various logistical problems encountered in sampling deep, rocky or reeflike
areas using either conventional surface-towed gear, or SCUBA.

In June 1979 a geophysical and biological survey of deep-water coral reefs
off Key Largo, Florida was conducted by the Office of Coastal Zone Man-
agement for NOAA, using a manned research submersible. Among the 55
specimens of decapod crustaceans collected were 2 new genera and 4 new
species. Included in the new taxa was a remarkable new genus of axiid
shrimp, described elsewhere (Kensley and Gore, 1980), an unusual new
genus of palaemonid shrimp, as well as the first western hemispheric record
for the rare Indo-West Pacific genus Pontoniopsis (Palaemonidae), and the
second western hemispheric record for the stenopodid shrimp genus Odon-
tozona. Other notable range extensions included the first western hemi-
spheric record for the alpheid shrimp Alpheopsis trispinosus (Stimpson),
and the first continental United States record for the pagurid crab genus

136 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Nematopaguroides Forest & St. Laurent, from Brazil. Additional conti-
nental U.S. records included specimens of Pseudocoutierea antillensis
Chace (Palaemonidae) and Synalpheus sanctithomae Coutiere (Alpheidae),
the former previously known from the unique type-specimen, and the latter
from a small type-series collected at various localities in the lesser Antilles.
The collection was thus remarkable both for the novelty of the contained
taxa, and from the fact that 11% of the 55 specimens, comprising 36 species,
were new to science.

Materials and Methods

The specimens in this report were collected from a series of 5 of 10
Stations occupied by the Research Submersible Johnson-Sea-Link I, oper-
ated by the Harbor Branch Foundation, Inc. Station data are listed in Table
1. In the following report, only new species, and new or interesting records
of previously known species are considered. The few remaining taxa in the
collections, all relatively common associates of coralline habitats, were of
less consequence and will not be discussed further, although they are listed
in Table 2. Catalog numbers for the species include those of the National
Museum of Natural History, Washington, D.C. (USNM) for new or excep-
tionally interesting taxa, and the Indian River Coastal Zone Museum
(IRCZM) at Link Port, Ft. Pierce, Florida. With the exception of holotypes,
most paratypes, and those species noted below in the text, the remainder
of the collection has been deposited in the Indian River reference collec-
tions.

Measurements were made with a substage micrometer calibrated to an
ocular reticle on a dissecting stereomicroscope, and the abbreviations are
as follows: rcl, rostral carapace length, measures the medial length from the
distal rostral tip to the posterior carapace margin; sl, shield length, was used
only with the pagurid species, and is the length of the carapace from the
rostrum posteromediad to the cervical groove.

The three new species in this study are named for my research assistants,
Mrs. Paula M. Mikkelsen, Miss Kim Allyn Wilson, and Mrs. Liberta E.
(Scotto) Poolt, in recognition of their cheerful and willing help both in the
field and in the laboratory over the past 8 years.

Section Caridea
Family Bresiliidae Calman, 1896
Discias atlanticus Gurney, 1939

Material examined.—1 °; rcl 3.2 mm; IRCZM 89:3802.

The single specimen constitutes the second record for the continental
United States, the species having been previously reported from the central
eastern Florida coast by Gore and Wilson (1978). Members of the genus are

137

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VOLUME 94, NUMBER 1

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138

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—List of decapod crustacean species collected during the deep reef survey Key
Largo Marine Sanctuary.

Family

Alpheidae

Bresiliidae
Hippolytidae

Palaemonidae

Processidae
Stenopodidae
Axiidae

Paguridae

Galatheidae

Xanthidae

Majidae

Genus/species

Alpheopsis trispinosus (Stimpson)

Alpheus amblyonyx Chace
Alpheus normanni Kingsley
Synalpheus pandionis Coutiere
Synalpheus sanctithomae Coutiere

Synalpheus townsendi Coutiere
Synalpheus sp. (?townsendi)

Discias atlanticus Gurney
Thor manningi Chace

Pontoniopsis paulae sp. nov.
Pseudocoutierea antillensis Chace
Veleroniopsis kimallynae n. gen. et sp.

Processa cf. tenuipes Manning & Chace
Odontozona libertae n. sp.
Coralaxius abelei Kensley & Gore

Nematopaguroides cf. fagei Forest &
St. Laurent

Munida angulata Benedict

Actaea bifrons Rathbun
Melybia thalamita Stimpson

Micropanope nuttingi (Rathbun)
Micropanope sculptipes Stimpson

Micropanope sp.

Juvenile xanthid

Pilumnus sp.

Pseudomedaeus distinctus (Rathbun)

Mithrax acuticornis Stimpson

Station

[-679
1-688
1-690
1-690
1-690
1-681
[-690
[-690
1-690

[1-679
[-682

[1-682
1-687
[-690

[1-679
1-688
[-679
1-690

[1-679
1-681
1-682
1-688
1-690

1-681
1-690
1-688
1-690
1-690
1-679
1-681
1-688
1-688
1-688
1-690
1-679

[1-679

Se el ee ee ole

Os +0 QO +10 0 BOW AA

No./Sex

2

66,228

2

3, 1 2 ovigerous

ovigerous

ovigerous

2 ovigerous

+O

366d, 1 2 (spent)
g

2 ovigerous

3

2 2 ovigerous

6, 1 2 ovigerous
Goll L

2 ovigerous
2
2
S51 25 1 TUN.
2° ovigerous
6,222

3 juv.
6,2 22, Il juny.

1 juv.
1 (crushed)

1

1

3 juv.
2 juv.

3

known associates with sponges, usually in water shallower than 40 m. The
76 m record in this study is the deepest yet for the species. Chace and
Brown (1978) discussed the status of and reason for synonymizing the family
Disciadidae with the Bresiltidae.

VOLUME 94, NUMBER | 3)

Family Palaemonidae Rafinesque, 1815
Subfamily Pontoniinae Kingsley, 1878
Pontoniopsis paulae, new species
Fig. 1A—P

Material examined.—Holotype, 1 3; rcl 3.2 mm; USNM 181240.

Diagnosis.—A small, short, heavy-bodied smooth pontoniine shrimp with
a short acute rostrum, and a massive major second cheliped bearing a dis-
tinct subdistal medial groove, which appears as if ventrally located when
cheliped is seen in lateral view.

Description.—Carapace smooth, shining, without grooves or ornamen-
tation; rostrum compressed, toothless, faintly carinate, acutely triangular in
dorsal view, produced into spinelike point, narrower than eye width, falling
short of distal margin of basal antennular segment; adrostral depressions
absent; orbital margin distinctly concave, shallowly excavate into frontal
margin, basal margin underlying ophthalmic peduncles. Lower margin of
orbit produced anteriorly into short, sharp, obscurely carinate antennal
Spine; no other carapacial spines present. Eyes noticeably ovoid, large dark
corneas. Basal antennular segment elongate, distolateral margin a spinous
projection extending to anterior margin of following segment, this and third
segment about equal in length and width; stylocerite very reduced, only
slightly produced laterally, rounded, covered by proximomesial expansion
of scaphocerite; latter lamellate, twice as long as wide, extending beyond
antennular peduncle, anterior margin convex, a distolateral spine overreach-
ing same; basal antennal peduncular segment unarmed, carpocerite similarly
so, latter narrowly cylindrical, about 5 times longer than wide, falling short
of distal end of basal segment of antennule. Mandibles without palp, molar
process with 4 blunt teeth, incisor with 7 bluntly rounded spines. Maxillule
and maxilla not dissected to avoid further damage to specimen. All maxil-
lipeds with well-developed exopods; maxilliped 1 with caridean lobe well
developed, bearing elongate lash, endopod entirely absent, a distinctly bi-
lobed, rectangular epipod; maxilliped 2 and 3 as illustrated, both with well-
developed epipod; maxilliped 3 barely attaining base of carpocerite and
antennular peduncle, a poorly developed arthrobranch present.

Pereopods | and 2 chelate. First pereopods equal, extending beyond distal
margin of antennular peduncle, chela reaching beyond scaphocerite; former
short, somewhat inflated, gaping, fingers 0.6 times longer than palm, distally
hirsute; carpus 0.8 times length of either chela or merus, about equal to
ischium, all segments smooth, unarmed. Second pereopods dissimilar, ex-
tending greatly beyond first pair. Major chela massive, smooth, unarmed,
inflated, elongate, sparsely setose; a distinct longitudinal groove subdorsally
on distal medial surface, opening into deep transverse notch paralleling an-
gle of fixed finger; fingers curved, gaping, shorter than palm, comprising
about % length of chela, equal to about half length of palm, a small rounded

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

140

VOLUME 94, NUMBER 1 141

tooth basally on cutting edge of each; 2 bluntly rounded teeth, plus several
small tubercles basally at exterior junction of dactyl and palm (Note: this
surface held ventrally when animal is alive, Fig. 1A); tips of both with small
tuft of hair; carpus short, smooth, shining, widely expanded distally, inner
distal angle a bluntly rounded tooth; merus 1.2 times longer than carpus,
proximally compressed, distally inflated, ventral surface flattened; ischium
compressed, shorter than merus, about equal in length to carpus. Minor
second pereopod shorter than major, chela elongate, conically cylindrical;
fingers curving laterally without noticeable gape, comprising about 0.4
length of chela, 34 length of palm, unarmed; carpus oblately pyramidal,
expanded distally, ventrally excavate along distal margin, 0.8 times length
of merus; latter cylindrical, equal in length to compressed ischium, all 3
segments unarmed. Pereopods 3, 4, 5, similar in configuration, slender, seg-
ments decreasing in length in relation to respective meri as follows: pro-
podus (0.9), carpus (0.6), dactyl (0.3); ventral margin of pereopod 3 pro-
podus with 3—4 spines, remaining legs with 3, other segments unarmed;
dactyls compressed, falcate, tips drawn into elongate, curved hooklike
spine, not bifid, but ventral margin with rectangular dentiform prominence
only slightly developed into supernumerary tooth. All pereopods apparently
lacking epipods.

Abdominal somites smooth, shining, unarmed; posterolateral margins of
somites 1-3 bluntly rounded, those of somites 4—5 more truncate; telson
about 1.4 times longer than somite 6, shorter than uropods, with 2 pairs
lateral spines placed just posterior to midpoint, and at posterior !/7 of length,
respectively; posterior margin with 3 pairs spines, intermediate pair about
4 times length of submedian, 6 times length of lateral pair at posterodistal
margin; uropods longer, narrower, than telson proper, distolateral angle of
exopod with strong, fixed tooth, followed by much smaller, movable spine;
a very faint diaresis present.

Gill formulae not determined to avoid further damage to specimen.

Ecology.—The single male was collected from the ventral surface of the
spatangoid echinoid Meoma ventricosa Lamarck, 1816, and is apparently
commensal with this species. The only other species in the genus, Ponto-
niopsis comanthi Borradaile, 1915 is a littoral form and a known commensal

<<

Fig. 1. Pontoniopsis paulae. Male, holotype. A, Lateral view; B, Cephalothorax, anterior
region; C, Same, lateral view; D, Antennule and antenna, ventral view; E, Mandible; F, Max-
illiped 1; G, Maxilliped 2; H, Maxilliped 3; I, Left pereopod 1; J, Pereopod 2, major (left)
chela, outer view (left), inner view (right); K, Pereopod 2, minor (right) chela, outer view; L,
Left pereopod 3, and detail of dactyl; M, Left pereopod 4; N, Left pereopod 5; O, Appendix
masculina, left pleopod 2; P, Tailfan, dorsal view. Scale lines as indicated.

142 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

with the crinoid Comanthus timorensis (Mueller) according to Borradaile
(1917). No crinoids were noted in the area where P. paulae, a decidedly
deeper water species, was collected.

Remarks.—Although agreeing in the generic diagnosis with characters
defining Pontoniopsis as provided by Borradaile (1917) and also delineated
in Holthuis’ (1952b) key, the Floridan species is not closely related to either
Pontoniopsis comanthi Borradaile, 1915, or to Pontoniopsis sp. noted by
Potts (1915) and apparently redescribed by Borradaile in his 1917 paper.
Holthuis (1952b) synonymized the latter with P. comanthi, although several
characters in Borradaile’s description differed from those described by Hol-
thuis from his specimen. Pontoniopsis paulae may be quickly distinguished
from P. comanthi by several important and easily observed features. The
‘most easily noticed of these include the distally-grooved major second che-
liped, the different relative lengths of the cheliped fingers to their respective
palms, the more spiniform rostrum, the lack of a postorbital marginal
groove, the position of the lateral spines on the telson and the different
lengths of the posterior submedian and intermediate pairs, and by the dis-
tolateral tooth of the scaphocerite which overreaches the anterior margin of
the lamella. These and other features are summarized in Table 3 where the
2 species may be easily compared.

The new species also shows some resemblance to Dasella herdmaniae
Lebour 1939, a pontoniine shrimp commensal with the ascidian Herdmania
in the Indian Ocean. In addition to the characters cited above, P. paulae
differs from D. herdmaniae in the armature of the dactyls on the walking
legs, in the absence of a palp on maxilliped 1, and in the much shorter
rostrum which does not extend beyond the very large eyes. Dr. A. J. Bruce
(in litt.) agreed that the two species bore a superficially close resemblance
to each other, but will demonstrate in a forthcoming paper still other dif-
ferences between the two taxa, emphasizing carapacial characters over-
looked by Lebour in her original description.

Distribution.—The genus Pontoniopsis was heretofore known from only
a few specimens recorded from the Indo-West Pacific (Holthuis, 1952b).
The discovery of the species described herein is the first time the genus has
been reported from the western North Atlantic. Presently it is known only
from the type-locality, Carysfort Reef, off Key Largo, Monroe County,
Florida (JSL I-682, Table 2).

Pseudocoutierea antillensis Chace, 1972

Material examined.—2 ° 2°, ovigerous; rcl 3.9, 3.6 mm; IRCZM 89:4666.
Both specimens agreed in every respect with the description given by
Chace, who also compared his new species with the eastern Pacific P.
elegans Holthuis, 1951, the only other member in the genus. Both species

VOLUME 94, NUMBER I

143

Table 3.—Comparison of morphological characters in two species of Pontoniopsis.

Rostrum:

Carapace:
Abdomen:

Telson:

Posterior
spines:

Stylocerite:

Scaphocerite:

Mandible:

Maxilliped 1:
Maxilliped 3:

Pereopod 1:

Pereopod 2:

Pereopods 3-5:

Pontoniopsis paulae

Narrow, spiniform

Smooth

Somites truncately rounded

1.4x longer than somite 6;
Spines placed 0.6, 0.8
length

Intermediate 4x larger than
submedial: 6x larger than
posterolateral

Greatly reduced

Extending beyond antennule
peduncular segments;
Distolateral spine reaches
beyond lamella margin

Molar with 4 processes,
incisor with 7 processes

No palp

Arthrobranch present;
appendage extending barely
to carpocerite base

Chela reaching beyond
scaphocerite; fingers longer
than palm; carpus shorter
than chela

Longer than pereopod 1;
fingers shorter than palm

Tips not bifid; obtuse
prominence present

Pontoniopsis comanthi [fide Holthuis, 1952b]

Lanceolate, compressed

Posterior margin or orbit with oblique
groove behind

Somites broadly rounded

Slightly longer than somite 6; Spines
placed 0.6, 0.8 length

Submedial greater than half length of
intermedial

Extending beyond middle of basal
segment

Extending slightly beyond antennule
peduncular segments

Distolateral spine falls short of lamella
margin

Molar with 4 processes, incisor with 3
processes

Palp present

Arthrobranch absent; appendage
extending well beyond carpocerite
base

Chela and carpus reaching beyond
scaphocerite; fingers equal to palm;
carpus longer than chela

Shorter than pereopod 1; fingers longer
than palm

Tips bifid; obtuse prominence not
mentioned.

are rare, but this may be a consequence of their habitat, rocky reeflike
regions with associated gorgonians and antipatharians, and thus an area not
easily sampled by conventional surface-towed gear. The 2 specimens from
Florida constitute the first continental United States record for the species,
which was previously known only from the type-locality, Saba Bank, north-
west of the Leeward Islands, approximately 1900 km southeast of Key

Largo.

144 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

VOLUME 94, NUMBER 1 145

Veleroniopsis, new genus
Pigs2A—P

Diagnosis.—A small, flattened pontoniine shrimp with a noticeably rect-
angular rostrum. Body dorsoventrally depressed overall, carapace smooth,
without spination of any kind; rostrum well developed, elongate, widely
expanded proximally, subrectangular distally, forming shelflike expansions
over eyes, with prominent median compressed, knifelike rostral spine; eyes
large, pigmented; scaphocerite well developed; mandibles without palp;
maxilliped 2 with exopod, maxilliped 3 lacking same, and without pleuro-
branch. Posterior orbital margin inflated, a noticeable postorbital tubercle;
anterior margin of carapace recurved to form tubelike opening into branchial
chamber. Pleura of first 5 abdominal somites more or less broadly rounded
or truncate, without ventral spines. Pereopods smooth, thin, elongate, dac-
tyli of last 3 pairs simple. Telson with 2 pairs of spines on posterior margin.

Type-species.—Veleroniopsis kimallynae, new species (see below).

Etymology.—A combination of Veleronia, a palaemonid genus to which
the new taxon shows some resemblance, and ‘‘opsis’’ from the Greek,
‘“‘*having the appearance of.’ Gender feminine.

Remarks.—The new genus bears much resemblance in a variety of char-
acters to several genera, including Lipkebe Chace, 1969, Veleronia Hol-
thuis, 1951, Anchistioides Paulson, 1875, Pontonides Borradaile, 1917, and
Neopontonides Holthuis, 1951. Veleroniopsis has, perhaps, its closest re-
lationships with the western Atlantic Lipkebe holthuisi and species of the
eastern Pacific Veleronia, especially in the notched anterior margin of the
carapace which forms a tubular opening into the branchial chamber, by the
rather obsolete or ill-defined posterolateral orbital margin, the third maxil-
liped lacking an exopod, by the broadly truncate rostrum expanded into
eaves above the orbits, and the simple dactyli of the last 3 pairs of pereo-
pods. The new genus differs from Veleronia in possessing a well-developed
lash on the first maxilliped (although this lash is also present in Lipkebe),
a second maxilliped with a well-developed exopod (absent in Lipkebe), and
in having a noticeable postorbital tubercle (in Lipkebe a spine is present at
this position). The dactyli of the walking legs in Lipkebe show a superficial

Fig. 2. Veleroniopsis kimallynae. Male, holotype. A, Lateral view; B, Frontal view; C,
Dorsal view; D, Antennule, dorsal view; E, Antenna, ventral view; F, Maxilliped 1; G, Max-
illiped 2; H, Maxilliped 3; I, Left pereopod 1; J, Left pereopod 2; K, Right pereopod 3; L,
Left pereopod 4; M, Left pereopod 5; N, Left pleopod 2 (exopod omitted); n, same, detail of
appendix masculina; O, Left pleopod 1, detail of endopod; P, Tailfan, dorsal view. Scale lines
as indicated.

146 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

resemblance to those in the new genus, but the chelipeds differ in relative
lengths of the fingers to the palms among the three genera discussed.

Although Veleroniopsis shares with the genus Anchistioides exopods on
maxillipeds 1 and 2, a postorbital tubercle, and 2 pairs (occasionally only
1 pair in the latter genus) of the telsonal marginal spines, the general mor-
phology of Anchistioides, such as the laterally compressed body and ros-
trum (the latter being toothed), the presence of an antennal spine, and the
minutely bifid dactyli on the last 3 pereopod pairs allows easy distinction.

Both Pontonides and Neopontonides resemble Veleroniopsis in some
features. However, in the former 2 genera the rostrum is either entirely
depressed without dorsal teeth, or is distally compressed, often with a vari-
able number of dorsal teeth, respectively. Pontonides is primarily an Indo-
West Pacific genus, exhibiting a distinct antennal spine, and rather large and
well-developed chelipeds, as opposed to no antennal spine, and elongate,
thin chelipeds in Veleroniopsis. Neopontonides, presently known from a
single species each in the eastern Pacific and western Atlantic, is easily
separated from Veleroniopsis by the form and dentition of the rostrum, as
noted above, by the presence of an endopod on the first maxilliped, and the
absence of an exopod on maxilliped 2. Moreover, both Pontonides and
Neopontonides carry 3 pairs of spines on the telsonal margin, thus sharing
this feature with Lipkebe and Veleronia, but not Anchistioides or the new
genus.

Two other genera showing a superficial resemblance to Veleroniopsis are
Coutierea Borradaile, 1917, and Pseudocoutierea Holthuis, 1951. Both of
the latter genera have well-developed antennal spines, the rostral expan-
sions are armed with either sharp supraocular teeth or elongate spines (sim-
ilar to that seen in Lipkebe), and the abdominal pleura of somites 3—5 are
drawn into sharp teeth, all features which are lacking in the new genus.

Veleroniopsis seems to be yet another genus with relationships toward
those pontoniine genera which have dispensed with, or are in the evolu-
tionary process of eliminating, exopods on the maxillipeds. It differs from
all of these in retaining (at least in the male) the exopod on the second
maxilliped. Several of these same genera share dorsoventrally depressed
carapaces, expanded or modified frontal or rostral regions, and in at least
2 instances having the anterolateral carapace margin modified into a tubelike
opening into the branchial chamber. It is, perhaps, also significant that these
genera have few species, usually 1 or 2, suggesting that the evolutionary
factors which produced this morphology, while successful to some degree,
have left little room for further expansion. The genera in question are also
known associates of coral reefs and reef-type habitats, including gorgonid
soft corals in some cases, all habitats where the ability to utilize small cracks
and crevices, or to lie more or less appressed against a substrate, may have
conferred selective advantage to the species against predation. Shaw et al.

VOLUME 94, NUMBER | 147

(1977), following an earlier suggestion by Bruce (1976) that the genus Lip-
kebe would probably prove to be a commensal, confirmed the relationship
with the crinoid Comactinia meridionalis meridionalis (Agassiz 1865). The
very close similarity between Veleroniopsis and Lipkebe suggests a similar
possibility, although no crinoids were noted in the habitat at the time the
new genus was collected.

Chace (1969) discussed the status of the various genera noted above,
establishing his new genus Lipkebe with some hesitation because the pre-
sumably generic characters used to define the other genera had resulted in
several monotypic taxa. Regrettably, Veleroniopsis becomes yet another
presently monotypic genus in this taxonomic grouping, although reconsi-
deration of perhaps the most important generic distinction, that of the pres-
ence of an exopod on maxilliped 2, could conceivably allow its placement
in Chace’s Lipkebe. Until further material becomes available, or the generic
groupings revised, I believe the establishment of the new genus to be war-
ranted.

Veleroniopsis kimallynae, new species

Material examined.—1 3; rcl 3.2 mm; USNM 181241. Holotype male.

Description.—Carapace elongate, smooth, shining, naked, microscopi-
cally punctate, dorsoventrally depressed; rostrum shelflike, broadly ex-
panded, flattened, elongate, excluding spine about '4 carapace length, wider
proximally, constricted mesially, bluntly rectangular distally, corners dis-
tinctly and broadly rounded, without spines; an elongate, acute, medial
rostral spine, appearing bladelike in lateral view, unarmed dorsally and ven-
trally, reaching just beyond scaphocerite distal margin, extending poste-
riorly onto expanded rostral portion as thin, carinate medial ridge, becoming
obsolete just posteriad to eyes; posterior orbital margins bluntly rounded,
a postorbital tubercle forming noticeable prominence viewed dorsally, ex-
tending backward from which an obsolescent ridge, anterior margin sweep-
ing obliquely forward and down past eyestalks to form thin plate, recurving
laterally outward and ventrally to form thin tubelike opening for branchial
chamber; lateral margin of latter very obscurely produced anteriorly into
blunt angle, extending posteriad as thin ridge, becoming obsolete at pos-
terolateral carapace margin; latter truncate, excavated dorsomedially; no
supraorbital, antennal, branchiostegal or pterygostomial spines.

Eyes enlarged, well-developed, protuberant, pigmented, stalks unarmed.
Antennular peduncle 3-segmented, barely reaching beyond tip of rostral
spine; basal article longest, about 3 times length of second, about twice
length of third, produced anterodistally into sharp tooth, mesially into sharp-
ly rounded small lobe, both extending to about midlength of following ar-
ticle; second article cylindrical, third truncate, both unarmed; upper flagel-
lum with 5—6 fused, 3—4 unfused articles, plus elongate hairlike whip; lower

148 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

flagellum thin, reaching beyond both pairs of extended chelae; stylocerite
reduced to small basal spine-tipped expansion on basal article. Antennal
peduncle 3-segmented; carpocerite proximal segment unarmed, distal seg-
ment 2.7 times longer than wide, slightly overreaching proximal margin of
antennular second article; scaphocerite unarmed basally, broadly lamellate,
produced distally into rounded lobe reaching about midlength of antennular
third article, greatly exceeding length of small, but distinct distolateral tooth;
first flagellar segment rounded, ball-like, remaining articles relatively stout,
sparsely setose, total length of flagella reaching region of second abdominal
somite.

Mandibles, maxillule and maxilla not dissected; former without palp.
Maxilliped 1 with well-developed bifid epipod, caridean lobe produced, a
thin lash; endopod wanting; maxilliped 2 with thin, well-developed epipod,
2-segmented exopod, 5-segmented endopod, ultimate segment of latter with
numerous sharp spines and stout setae; maxilliped 3 with epipod, lacking
exopod; endopod 3-segmented, meral and carpal articles fused, ultimate
segment with spines and stout setae as in maxilliped 2.

Pereopod | thin, chelate, reaching when extended beyond antennular pe-
duncle; ischium truncate, slightly inflated distally, unarmed; merus about
twice ischial length, smooth; carpus distally expanded, shorter than merus,
unarmed; chela subequal to merus, fingers about equal to palmar length,
gaping, upper crossing over lower, gape of movable finger with thin irregular
tooth about midway, tips of both fingers with tuft of hairs and elongate,
crooklike setae; except for sparsely scattered hairs cheliped naked. Pereo-
pod 2 more robust than former, chelate, elongate, extending well beyond
antennular peduncle by most of chelae; ischium elongate, smooth, unarmed;
merus slightly shorter (about */6 length) of ischium, barely inflated distally;
carpus short, curved, slightly less than half meral length; chela heavy, elon-
gate, about 3.5 times longer than carpus, fingers equal to palmar length,
hooked at tip, each bearing small apical tuft of setae, gaping slightly but
evenly, unarmed. Pereopods 3-5 similar in morphology to each other,
smooth, unarmed, sparsely setose except distal margin of propodi which
carry tufts of elongate setae and fine hairs, dactyli bluntly falcate, simple,
without supernumerary teeth.

Sternal plate wide, flattened, smooth, shining. Abdominal somites dor-
soventrally depressed, smooth, broadly and evenly rounded dorsally, very
sparsely setose, unarmed; pleura truncately rounded, posterolateral angles
on somite 3 bluntly angular, that of somites 4, 5, ellipsoidal; somite 6 about
2.3 times longer than fifth, a faint dorsomedial carina. Pleopods on somites
1-5, peduncles unarmed; rami elongate, foliaceous, first 2 pairs when ex-
tended forward reaching level of coxae of pereopod 3; pleopod 1 endopod
modified into shortened, bluntly spatulate accessory gonopod bearing sev-
eral setae basally; pleopod 2 with setose appendix masculina; endopods

VOLUME 94, NUMBER | 149

shorter than exopods on all five pairs, those of second to fifth pairs with
well-developed appendices internae.

Telson about 1.2 times as long as somite 6, narrow, rounded dorsally, 2
pairs of minute lateral spines on margins of posterior half, placed about 74
and °/6 total telson length, respectively; posterior margin with 2 pairs spines,
inner pair about 4 length of outer, plus 4 elongate setae on posterodistal
margins; uropodal exopod and endopod about equal in length, former more
expanded, with distinct fixed spine at distolateral angle; endopod elongate,
narrow, margins without spines; both appendages heavily setose.

Remarks.—As noted in the discussion under the generic description, Ve-
leroniopsis kimallynae is an unusual palaemonid shrimp. Although appear-
ing similar to the western Atlantic Lipkebe holthuisi Chace, and to species
of the eastern Pacific genus Veleronia, it is easily distinguished from both
of these by the rostral configuration, the telsonal spine and setal formula,
and the exopod on the second maxilliped. The appendix masculina illustra-
tions of Lipkebe holthuisi provided by Shaw et al. (1977) are superficially
similar to that of V. kimallynae, but differ in lateral spination being present
on the former species, but absent in the latter. Unfortunately, the female of
V. kimallynae is as yet unknown so any variation or sexual dimorphism
within the species remains to be determined.

The general habitat, relict Montastraea coral, may explain why the
species has not been discovered until now, for unless such large and pon-
derous coral heads are carefully pulverized and the debris examined in
sieves, many inquiline species will be missed.

Distribution.—Known only from the type-locality, Elbow Reef, off Key
Largo, Monroe County, Florida (JSL I-690, Table 2).

Family Alpheidae Rafinesque, 1815
Alpheopsis trispinosus (Stimpson, 1861)
Fig. 3A—G

Material examined.—2 66,2 22 (1 ovigerous); rcl 3.8, 3.6; 5.1, 2.3 mm;
IRCZM 89:4667; 1 6, 89:4668; 1 2° ovigerous, gift to A. H. Banner.

The present material, with distinctly trispinose front, the carapace lacking
any dorsal carinae, and both chelae bearing a distinct longitudinal groove,
differs from all known western Atlantic species of Alpheopsis, although it
is extremely close to A. africanus Holthuis, 1952a. The material was sent
to Dr. and Mrs. A. H. Banner for confirmation, and they identified it as
Stimpson’s species, pointing out that the chelipeds in the Florida specimens
were equal both in size and length, whereas those appendages in A. afri-
canus are distinctly unequal. The Floridan specimens agreed in all important
respects with the redescription and subsequent neotype designation for A.
trispinosus (Banner and Banner, 1966), differing only in relative proportions

150 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Alpheopsis trispinosus. Male, IRCZM 89:4668. A, Cephalothorax, anterior region;
B, Left major chela, lateral view; C, Same, dorsal view; D, Left pereopod 2; E, Right pereopod
3; F, Same, detail of distal articles; G, Tailfan, dorsal view. Scale lines equal 1.0 mm.

VOLUME 94, NUMBER I 151

of rostral and frontal spines, carpal articles of pereopod 2, dactylar length
on pereopods 3-5, and telsonal length-width ratio. None of these differences
are, in my opinion, sufficiently significant to warrant establishment of a
western Atlantic subspecies at this time. I thus concur with the suggestion
made by Mrs. Banner (in /itt.) that Alpheopsis trispinosus may be pantrop-
ical in distribution. Previous records in the literature are from the Australian
region, with possible distribution in the tropical Pacific and perhaps in the
eastern Atlantic off the Azores, as well (see Banner and Banner 1966).

Five other specimens (1 6,3 2 2, ovigerous) collected from off the central
eastern Florida coast by R/V Gosnold (G-222/274D; 27°52.5'N, 79°57.5'W,
75-98 m, Smith-MacIntyre grab, 28 February 1974.—G 246/709; 27°44.3'N,
79°58.0'W, 72 m, pipe dredge, 4 September 1974) possess a trispinose front
and lack carapacial carinae. But unfortunately, all are mising chelipeds so
specific identification remains tentative.

Alpheus amblyonyx Chace, 1972

Material examined.—1 °; rcl 2.9 mm; IRCZM 89:3791.

Previously recorded from Dominica, Puerto Rico, St. Thomas, Bahia de
la Ascension, Quintana Roo, and the northwest Gulf of Mexico (Chace,
1972; Pequegnat and Ray, 1974), the Key Largo female is the second to be
recorded from the continental United States. However, several other spec-
imens of this species, as weil as some which appear to be more closely
allied to Alpheus macrocheles Hailstone, 1835, have been collected from
offshore stations on the central eastern Florida coast by R/V Gosnold. The
species range can thus be considered as extending from the lesser Antilles
to eastern Florida. Depth range is from about | to 24 m. Some Indian River
region material off central Florida collected by R/V Gosnold in 70-90 m,
plus a single cheliped from 135-145 m, would extend the depth range con-
siderably if all this material belongs to A. amblyonyx.

Synalpheus pandionis Coutiere, 1909

Material examined.—1 ¢; rcl 3.9 mm; IRCZM 89:3789.

The species is rare in collections and was previously known from the
eastern and western Gulf of Mexico, and the Virgin Islands southward to
Barbados and Curacao. Undoubtedly a coral associated species, both the
Key Largo specimen, and a spent female from off St. Lucie Inlet on the
central eastern Florida coast, were taken from relict coral heads. The male
was taken in 18 m, the female in 48 m, well within the recorded depth range
from the intertidal to 80 m (Chace, 1972; Pequegnat and Ray, 1974).

152 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Synalpheus sanctithomae Coutiere, 1909

Material examined.—1 3, 1 2 ovigerous; rcl 3.9, 3.8 mm, respectively;
IRCZM 89:3799, 89:3792.

The identification of the 2 specimens, which conformed quite well with
Coutiere’s (1909) original account and illustrations, was confirmed by Dr.
R. B. Manning who compared the Florida material with Coutiere’s syntypic
series. Until now, the species was known from only 3 specimens, including
the two designated as types by Coutiere, collected off St. Thomas. The Key
Largo material extends the range approximately 1800 km northwestward
from the type locality, and constitutes the first continental United States
record for the species. Although the color in life was not noted, the fingers
on the major chela of the male from Key Largo are a distinct golden yellow
after more than one year in preservative.

Family Hippolytidae Bate, 1888
Thor manningi Chace, 1972

Material examined.—1 @ ovigerous; rcl 2.2 mm; IRCZM 89:3795.

This widely distributed little grass shrimp has been found associated with
intertidal grassflats, and subtidally (previously to 11 m) with living and dead
coral. The single specimen from 62.5 m on Carysfort Reef, among algal
covered rock, is the deepest yet recorded.

Family Processidae Ortmann, 1896
Processa cf. tenuipes Manning & Chace, 1971

Material examined.—1 °; rcl 5.9 mm; IRCZM 89:4669.

Although exhibiting general characters which allow it to be placed near
P. tenuipes in Manning and Chace’s (1971) key, the specimen differs from
that species in having the distal interior angle of the stylocerite drawn into
a distinct spine instead of bluntly angled, in having 12 and 33 articles in the
merus and carpus of the second leg (instead of 5-9, 17-26, respectively),
and in lacking the characteristic transverse row of fine hairs across the
dorsal base of the telson. The specimen was compared with type-material
in the National Museum of Natural History by Dr. R. B. Manning, who
informed me (in litt.) that he suspects it to be undescribed. The single
specimen is, unfortunately, incomplete, lacking the right pereopod 2, and
several other walking legs, so establishing it as a new species must await
additional material.

The Key Largo female was collected from coralline rock on a sandy
substratum in 42-76 m. Processa tenuipes also appears to prefer deeper
waters (31—331 m) and has been taken off North Carolina, in the northeast-
ern Gulf of Mexico, and a single specimen from off Havana, Cuba. All were
associated with fine sands or clay oozes.

VOLUME 94, NUMBER | 153

Section Stenopodidea
Family Stenopodidae Huxley, 1878
Odontozona libertae, new species

Figs. 4A—C, 5A-L

Material examined.—1 3, Holotype, rcl 3.6, USNM 181242; 1 @, Allo-
type, rcl 3.2, USNM 181243; 1 3, paratype, rcl 2.8 mm (rostrum broken);
USNM 181244.

Diagnosis.—A small, robust, heavy-clawed, spiny shrimp, with a distinct
cincture of spinules along posterior margin of cervical groove, smooth ab-
dominal somites, a heavily spined manus on pereopod 3, maxilliped 3 is-
chium with a ventrolateral row of spinules, and distinctly biunguiculate dac-
tyls on the walking legs.

Description.—Rostrum compressed, bladelike, slender, straight, about 34
or less carapace length, reaching distal margin of antennular second article,
falling far short of scaphocerite tip; armed dorsally with 5, ventrally with
45 spines. Carapace short, robust, heavy, microscopically punctate; gastric
region inflated, 2 pairs large medial spines about center and on posterior
slope, 2—3 others of varying size on anterior slope above orbital margin; a
smaller postorbital spinule; cervical groove distinct, postcervical groove less
prominent, posterior margin of former with cincture of about 14 large, hor-
izontal, anteriorly-directed spines; latter groove with interrupted transverse
dorsal rows of about 8—10 smaller spinules, between which lie 2 larger
spines, plus lateral row of 12-16 spinules of decreasing size, extending
obliquely anteriad; dorsal and lateral carapace surface between these 2
grooves with scattered, short, forwardly-directed spinules. Posterodorsal
margin concave, unarmed medially, a single row of 10-12 tiny spinules
dorsolaterally, preceded by shorter, similar row of about 6; ventrolateral
margins oblique, unarmed. Antennal spine large, massive, branchiostegal
spine smaller, both with faint carina; pterygostomial spine smallest, may be
doubled, posterior to which | or 2 larger spines parallel anteroventral mar-
gin; branchial ridge present, becoming more or less obsolete posteriad. Or-
bital margins formed medially from rostral base, becoming obsolete laterad;
eyes large, corneas inflated, pigmented, an interrupted series of several
small spinules along corneal base dorsally and anteriorly.

Antennular peduncle 3-segmented; basal segment elongate, exceeding
length of following 2 articles, somewhat sculptured, curved dorsally, 1 or
more small spinules on distal dorsal margin; second segment about 4% length
of first, third still smaller, both unarmed; flagella greatly elongate, whiplike,
extending beyond telson when complete, articles noticeably setose. Anten-
nal peduncle with 3 large segments, basal and penultimate largest, latter
subglobose, 2 proximoventral, 2 distoventral spinules; carpocerite about 4
length of scaphocerite, unarmed; scaphocerite blade narrow, elongate, la-
mellate, far overreaching rostral spine tip, 4 times longer than wide, spi-

154 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

4
Jt ee
ae

ber

Fig. 4. Odontozona libertae. Male, holotype. A, Dorsal view; B, Lateral view, pereopods
not illustrated; C, Female allotype, lateral view, pereopods not illustrated. Male right pereopod
3 and chela broken. Scale lines equal 1 mm.

VOLUME 94, NUMBER 1 155

nulate along concave outer margin, a strong apical spine, heavily setose
along convex inner margin, dorsal and ventral surfaces unarmed; flagella
extremely long, whiplike, reaching to or beyond telson, articles noticeably
setose.

Mandibles heavily chitinized, scoop-shaped molar process bluntly round-
ed, an oblique dentate protuberance, incisor process 2 strong blunt teeth;
3-segmented palp, setose along distal margin of second article, heavily
spined over surface of third. Maxillule and maxilla as illustrated, both with
well-developed endopod; coxal and basal endites distinct; scaphognathite
heavily setose. Maxillipeds 1-3 with exopods; first maxilliped with large,
bifid epipod, a distinctly setose endopod; second maxilliped endopod 5-seg-
mented, a noticeable arthrobranch and simple epipod; third maxilliped en-
dopod extremely elongate, reaching beyond scaphocerite tip by distal half
of propodal, and entire dactylar segment, all except last article armed ven-
trally or ventrolaterally with series of small spinules; mesially a series of
elongate, strong setae mesh with those on opposite appendage, plus several
still longer setae at dactylar-propodal junction; exopod long, multiarticulate
distally, attaining midlength of meral article; epipod simple.

First 3 pereopods chelate, elongate, increasing in size posteriad, all reach-
ing to, or well beyond, scaphocerite tip; each with epipod. Pereopod | thin,
smooth, without spines except for 2 distoventrally on carpus; latter article
longest, ischium about 0.6 times length of same, 0.8 times length of merus;
merus about %4 length of carpus, inflated distally; chela about %4 carpal
length, compressed, distally hirsute, unarmed; fingers gaping, crossing at
tips, with series of small, more or less evenly rounded or serrate teeth along
length, becoming irregular distally; movable finger half length of palm. Per-
eopod 2 similar to first in shape, larger, more robust, unarmed; articles
increasing in length from ischium to carpus, former about half carpal length,
274 meral length; merus about 0.7 times carpus, inflated distally, without
spines; chela about %4 carpal length, compressed, bearing several separated
tufts of long, distally curved setae, plus dense tuft of hairs apically; fingers
gaping, with irregularly spaced, ragged teeth on cutting edges, crossing at
tips, about 0.4 times length of palm. Pereopod 3 largest, most robust, articles
increasing in length and armature distally; ischium about half length of chela,
a single dorsodistal spine; merus about 0.6 times chela length, an interrupted
series of short, curved spines along dorsal and ventral margins, strongest
at distal angles; carpus about 0.8 times chela length, a paired series of 8-9
spines dorsally, becoming larger, more curved distally, a second series of
single spines along ventral margin, enlarging as in preceding series; manus
interiorly smooth, with 2 irregular paired rows of sharp spines along dorsal
margin, a smaller spinule on mesioventral margin below propodal-dactylar
junction; fingers thick, heavy, crossing at tips, angled slightly inward from
plane of manus, about 0.7 times length of palm, slightly gaping, armed along

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

156

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VOLUME 94, NUMBER | 157

cutting edges with irregular series of minute spinules; movable finger with
2 small, widely-spaced mediodorsal spinules, plus single triangular basal
tooth fitting into similarly shaped excavation on fixed finger. Pereopods 4
and 5 thin, fifth longest, carpi and propodi on both multiarticulate, dactyli
distinctly biunguiculate. Pereopod 4 carpus longest, 8-segmented, length of
segments numbered distally as: 5=7=8, 4, 6, 3, 2, 1, all unarmed; propodus
about half carpal length, 5-segmented, lengths numbered distally increasing
as: 2, 1, 4, 3, 5, all armed ventrally with short spinules, numbered from first
to last 2, 4, 4, 4-5, 6-8; merus 0.7 times, ischium about 0.4 times, carpal
length, both unarmed; a simple epipod. Pereopod 5 similar to preceding,
carpus longest, eighth article indistinctly separated, segments increasing in
lengths as 5, 8=7, 2, 6, 4, 3, 1, all unarmed; propodus about half carpal
length, 6-segmented, lengths increasing as: 2, 5, 1=4, 3, 6, with ventral
series of spinules progressing distally, 2, 3, 5, 5, 3, 7; ischium about 4,
merus about % carpal length, both unarmed; no epipod.

Dorsal and lateral surfaces of abdominal somites without spines or spi-
nules; first to third each with transverse grooves and carinae along anterior
dorsal margin, sculptured so that posterior part of somite 2 and 3 raised up
and overriding anterior carinae of same somites; third somite largest, pos-
terior margin overlapping anterior margin of fourth, latter without transverse
grooves or Carinae; somite 5 smallest, smooth; somite 6 with single oblique
row of small spinules on dorsolateral surface. Ventral and posterior margins
of pleura in males more spinose than females, pleuron of somite 1 large,
acute, directed ventrally, followed by smaller tooth at posteroventral mar-
gin; somites 2—4 with distinct spinous tooth at antero- and posteroventral
angles, pleuron of somite 4 also produced ventromedially into tooth, plus
2 smaller spines dorsad to posteroventral angle; somite 5 posteroventral
angle with teeth more noticeably spiny; somite 6 ventrolateral angle pro-
duced into distinct tooth, a single lateral spine, plus a second anterolateral
spinule just beneath posterolateral series of preceding somite. Abdominal
pleonal surface in male each with single median spine. Female abdominal
sculpturing and armature on pleura more or less similar to that of male but
much less developed; spines on abdominal pleonal surface lacking.

Pleopods on somites 1-5, those of first uniramous, remaining biramous;

<<

Fig. 5. Odontozona libertae. Male, paratype. A, Left mandible, sternal view; a, Same,
interior view; B, Maxillule; C, Maxilla; D, Left maxilliped 1, sternal view (left), interior view
(right); E. Left maxilliped 2, sternal view (left), interior view (right); F, Maxilliped 3; f, Same,
detail of meral article; G, Left pereopod 1; g, Same, detail of chela; H, Left pereopod 2; h,
Same, detail of chela; I, Left pereopod 3; J, Left pereopod 4; K, Left pereopod 5; L, Pleopod
2, female allotype (left), male paratype (right). Scale lines equal 1.0 mm.

158 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

peduncles of males triangular in cross-section, serrated posteriorly, a small,
short, sharp spinule basal to endopod; peduncles of females more lamellate,
non-serrate, lacking spinule, but with row of straight thin setae; no appendix
masculina; both sexes without appendices internae.

Uropodal peduncle produced distally into large sharp spine; dorsal surface
of exopod with 2 longitudinal, unarmed carinae; lateral margin bearing 4—6
teeth, including largest at distolateral angle; endopods with single unarmed
median carina, 2 lateral spines proximal half of appendage. Telson elongate,
expanded proximally, narrowing distally, 2 longitudinal carinae on dorsal
surface, separated by deep longitudinal groove; dorsal surface armed with
16 large, strong spines as follows: basally, one each laterally, plus a pair
medially; these followed distally by 4 spines, paired on each carina (occa-
sionally an extra spine as in holotype male), last pair of spines terminating
each carina; 2 strongest spines on telson lateral margin about midlength, a
smaller spine at each posterolateral angle. Uropodal and telsonal margins
heavily clothed with long setae; individual longer, hairlike setae dorsally on
same articles, including triple setae at base of uropodal peduncular spine.

Ecology.—The 3 specimens on which the preceding description was based
were all collected from a sponge and Siliquaria (Mollusca) colony. It seems
probable, based on the structure of the chelipeds, and the greatly lengthened
antennular and antennal flagella, that the genus and species is a “‘cleaning
shrimp.’’ Other members of the family, most notably the genus Stenopus,
are well known for their symbiotic relationships with various species of reef-
dwelling fishes, attracting attention of their hosts by waving and snapping
their brightly colored flagella and rocking slowly from side to side. Regret-
tably, no behavioral observations were made when QOdontozona libertae
was collected so the exact behavioral sequence remains as yet unknown for
this species.

Remarks.—The genus was previously known from West Africa and the
Indo-West Pacific region (e.g. Holthuis, 1946) and it was only recently dis-
covered to occur in the western Atlantic (Goy, in press). With the finding
of the second species described herein, and thus constituting the first record
for the continental United States, the genus may safely be considered to be
established in the tropical western Atlantic.

Odontozona libertae differs from O. spongicola (Alcock and Anderson,
1899) in possessing spinules behind the cincture of the cervical groove, these
being absent in the latter species. The new species differs from Odontozona
sculpticaudata Holthuis, 1946, in the third abdominal somite, which lacks
distinct longitudinal grooves, and in the absence of transverse grooves on
somites 4-6. The carapace is also much less spinulose in the new species
than in O. sculpticaudata. Although superficially resembling Odontozona
ensifera (Dana, 1852) the new species is easily distinguished by the general
armature on the abdominal pleura, by the lack of a transverse carina on

VOLUME 94, NUMBER 1 159

abdominal somite 3, by the more poorly armed rostrum, by the greatly
reduced number of spinules on the postcervical groove, fewer paired spines
on the telsonal carinae, the longer basal antennular segment, and by the
scaphocerite greatly overreaching the tip of the rostral spine. The descrip-
tion of Odontozona edwardsi (Bouvier, 1908) is incomplete (fide Holthuis,
1946), but O. libertae is easily separated from this species by a greater
number of segments in the carpus and propodus of pereopods 4 and 5 (8,
5; 8, 6; as opposed to 4, 5; 4, 5, in O. edwardsi). Odontozona libertae will
be seen to differ from Goy’s new species (in press) primarily by the less
developed spinulation on the carapace of O. libertae, by the less sculptured
abdominal somites, the distinctly spinose third chela, and a telsonal spine
formula differing from Goy’s material.

Distribution.—At present the species is recorded only from the type-lo-
cality, Elbow Reef, off Key Largo, Monroe County, Florida (JSL 1-688,
Table 2). The genus can be considered to be more or less pantropical, al-
though it is not yet known from the eastern Pacific region.

Section Thalassinidea
Family Axiidae Huxley, 1879
Coralaxius abelei Kensley & Gore, 1980

Material examined.—1 2; rcl 4.5 mm; USNM 173629.

This species, described from the female collected off Key Largo, and
additional material from Carrie Bow Cay, Belize, has been previously con-
sidered by Kensley and Gore (1980).

Section Anomura
Family Paguridae Latreille, 1803
Nematopaguroides cf. fagei Forest & St. Laurent, 1967

Material examined.—1 ¢ juv; sl 1.1 mm; IRCZM 89:4699.

Forest and St. Laurent’s species was described from only 3 specimens,
the male holotype and 2 very small (ca. 2 mm sl) females, one of which was
ovigerous. If the Key Largo specimen is indeed this species, then it would
appear that N. fagei is a remarkably small pagurid. The single specimen
was collected from a habitat similar to that occupied by those originally
described from Brazil, viz. coralline rock. Dr. Patsy A. McLaughlin who
examined the Key Largo juvenile stated that it appeared to belong to N.
fagei, but because of the small size positive identification could not be made.
She noted that the Floridan specimen differed from the description of N.
fagei in having fewer spines on the dorsal surface of the left cheliped carpus,
the dactyl of the right cheliped lacked the minute tubercles described in the
type from Brazil, and the telson was slightly different from that illustrated
by Forest and St. Laurent (1967). The Floridan material, if correctly as-

160 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

signed, constitutes a range extension of approximately 6300 km northward
from the type locality, off Recife, Brazil.

Family Galatheidae Samouelle, 1819
Munida angulata Benedict, 1902

Material examined.—3 33,6 2 2 (5 ovigerous); rcl 4.6—5.0; 3.2-5.9 mm;
IRCZM 89:3788, -3790, -3797, -3879, -3801.

Mrs. Barbara Shuler Mayo had previously informed me (in Jitt.) that
Benedict’s (1902) illustrations of M. angulata (Fig. 4, p. 253) and M. media
(Fig. 12, p. 263) have been transposed, so that of M. media actually illus-
trates M. angulata, and vice versa. The oblique angle of the cheliped, to
which the specific epithet angulata refers, is clearly seen in Benedict’s Fig.
12. Munida angulata has been recorded from the Gulf of Mexico (Benedict,
1902; Pequegnat and Ray, 1974) and off the central eastern Florida coast
(Avent et al., 1977). The Key Largo records suggest that the species will be
found around the southern tip of the Floridan peninsula.

Discussion

It is not often that a collection as small as the one made available contains
such a rewarding number of new genera, species, and records. This is es-
pecially true because the specimens were obtained in an area previously
considered to be rather well known in regard to the decapod crustacean
fauna. If nothing else the Key Largo collections underscore the caution that
must be exercised when statements concerning the knowledge of such a
fauna are made. The poorly sampled areas, such as the deeper, rocky and
coralline reef areas, remain for the most part terra incognita, especially for
the associated cryptic fauna inhabiting the crevices within relict or living
coral heads. It may well be that further knowledge of this fauna can only
be gained by in situ sampling using manned submersibles or lock-out divers,
because of the attrition of nets and trawls towed from the surface in such
topographically complex regions.

Literature Cited

Alcock A., and A. R. Anderson. 1899. An account of the deep-sea Crustacea dredged during
the surveying-season of 1897-98. Natural history notes from H.M. Royal Indian Marine
Survey Ship “‘Investigator**, Commander T. H. Heming, R.N., commanding.—Series
III, No. 2.—Ann. Mag. Nat. Hist. ser. 7, 3:1-27, 278-292.

Avent, R. M., King, M. E., and R. H. Gore. 1977. Topographic and faunal studies of shelf-
edge prominences off the central eastern Florida coast.—Int. Revue Ges. Hydrobiol.
62(2): 185-208.

Banner, A. H., and D. M. Banner. 1966. The alpheid shrimp of Thailand.—The Siam Society,
Monograph series, No. 3:i-iv, 1-168.

VOLUME 94, NUMBER | 161

Benedict, J. E. 1902. Description of a new genus and forty-six new species of crustaceans of
the family Galatheidae, with a list of the known marine species.—Proc. U.S. Nat. Mus.
26:243-334.

Borradaile, L. A. 1915. Notes on Carides.—Ann. Mag. Nat. Hist. ser. 8, 15:205-213.

. 1917. On the Pontoniinae. The Percy Sladden Trust Expedition to the Indian Ocean

in 1905, under the leadership of Mr. J. Stanley Gardiner.—Trans. Linn. Soc. Lond.,

Zool. ser. 2, 17:323—396.

Bouvier, E. L. 1908. Sur les relations zoologiques des Crevettes de la tribu des Stenopides.—
C. R. Acad. Sci. Paris 146:887-891.

Bruce, A. J. 1976. A further occurrence of Lipkebe holthuisi Chace (Decapoda Natantia,
Pontoniinae).—Crustaceana 30(3):310—312.

Chace, F. A., Jr. 1969. A new genus and five new species of shrimps (Decapoda, Palaemon-

idae, Pontoniinae) from the western Atlantic.—Crustaceana 16(3):251-—272.

. 1972. The shrimps of the Smithsonian-Bredin Caribbean Expeditions with a summary

of the West Indian shallow-water species (Crustacea: Decapoda: Natantia).—Smithson-

ian Contrib. Zool. 98:i-x, 1-179.

, and D. E. Brown. 1978. A new polychelate shrimp from the Great Barrier Reef of

Australia and its bearing on the family Bresiliidae (Crustacea: Decapoda: Caridea).—

Proc. Biol. Soc. Wash. 91:756—766.

Coutiere, H. 1909. The American species of snapping shrimps of the genus Synalpheus.—
Proc. U.S. Nat. Mus. 36(1659):1—93.

Dana, J. D. 1852. Crustacea, Part I—United States Exploring Expedition during the years

1838, 1839, 1840, 1841, 1842 .. . Vol. 13:i—viii, 1-685.

. 1855. Ibid. Atlas:1—127, pls. 1-96.

Forest, J., and M. de Saint Laurent. 1967. Crustacés Décapodes: Pagurides.—Result. Scient.
Camp. Calypso. Fasc. 8(6):47-169.

Gore, R. H., and K. A. Wilson. 1978. Studies on decapod crustacea from the Indian River
region of Florida, X. A first continental record for Discias atlanticus Gurney, 1939
(Caridea, Disciadidae).—Crustaceana 35:109-111.

Gurney, R. 1939. A new species of the decapod genus Discias Rathbun from Bermuda.—
Ann. Mag. Nat. Hist. ser. 11, 3:388-393.

Hailstone, S. 1835. Descriptions of some species of crustaceous animals; with illustrations
and remarks by J. O. Westwood.—Mag. Nat. Hist. 8:261-277, 394, 395, 549-553.
Holthuis, L. B. 1946. The Decapoda Macrura of the Snellius Expedition. I. The Stenopodidae,
Nephropsidae, Scyllaridae and Palinuridae. Jn Biol. Result. Snellius Exped. XIV.—

Temminckia 7:1-178.

. 1951. A general revision of the Palaemonidae (Crustacea Decapoda Natantia) of the

Americas. II. The subfamilies Euryrhynchinae and Pontoniinae.—Allan Hancock

Found. Occas. Pap. 11:1—332.

. 1952a. Crustacés Décapodes, Macrures.—Expéd. Océanog. Belge dans les Eaux Co-

tieres Africaines de |’ Atlantique Sud (1948-1949). Résult. Sscient. 3(Fasc. 2):1-88.

. 1952b. The Palaemonidae collected by the Siboga and Snellius Expeditions with re-

marks on other species. II. Subfamily Pontoniinae.—Siboga Expeditie XX XIXa!°: 1-253.

Kensley, B., and R. H. Gore. 1980. Coralaxius abelei, genus et species novum (Crustacea
Decapoda: Thalassinidea; Axiidae) a remarkable coral-inhabiting shrimp from the Flor-
ida Keys and the western Caribbean Sea.—Proc. Biol. Soc. Wash. (in press).

Manning, R. B., and F. A. Chace, Jr. 1971. Shrimps of the family Processidae from the
northwestern Atlantic Ocean (Crustacea: Decapoda: Caridea).—Smithsonian Contrib.
Zool. 89:1—41.

Paulson, O. 1875. Investigations on the Crustacea from the Red Sea with notes on Crustacea

162 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

of the adjacent seas. Part 1. Podophthalmata and Edriophthalmata (Cumacea):i—xiv, 1—
144 [Translation: Israel Program for Scientific Translations, 1961; National Science
Foundation and Smithsonian Institution, Washington, D.C.].

Pequegnat, L. H., and J. P. Ray. 1974. Crustaceans and other arthropods. Jn Biota of the
West Flower Garden Bank, Pt. I1.—Publ. Flower Garden Ocean Res. Cent.:232-288.

Potts, F. A. 1915. The fauna associated with the crinoids of a tropical coral reef: with special
reference to its color variation.—Carnegie Inst. Wash. Pap. Dept. Mar. Biol. III:73—97.

Shaw, J. K., Heard, R. W., Jr., and T. S. Hopkins. 1977. Notes on the biology of the
pontoniine shrimp Lipkebe holthuisi Chace, with a description of the male.—Proc. Biol.
Soc. Wash. 90:284—290.

Stimpson, W. 1861. Prodromus descriptionis animalium evertebratorum quae in expeditione
ad Oceanum Pacificum Septentrionalem ... Pars VIII. Crustacea Macrura.—Proc.
Acad. Nat. Sci. Philadelphia. pp. 24—46 [Proc. for 1860].

Smithsonian Institution, Ft. Pierce Bureau, Ft. Pierce, Florida 33450.

Note added in proof: In the course of this study the manuscript and
illustrations describing the 3 new species were sent to Drs. L. B. Holthuis
and A. J. Bruce to obtain their advice. Because of delays, replies were
not received until after the final manuscript was accepted for publication.
According to Dr. Bruce, the new species of Pontoniopsis does not much
resemble members of that genus, and only shows superficial resemblance
to Dasella. Dr. Holthuis agreed but also thought it not close to Dasella,
but rather to Onycocaris. Using Holthuis’ (1955) key, the characters could
conceivably place the new species either in Pontoniopsis or Onycocaris,
based on rostral configurations. It does not, however, agree with any
known species within the latter genus, using Bruce’s key (J. nat. Hist.,
1971, vol. 5, pp. 293-298). It is apparent, especially when Bruce’s work
with Indo-Pacific pontoniine shrimps is considered, that many species show
intermediate relationships within established genera, often requiring (as
Bruce has done) establishment of new genera. This may eventually prove
to be the case with Pontoniopsis paulae. Until additional material becomes
available, and the limits of the pontoniine genera to which Pontoniopsis
paulae shows affinity are better defined, I have assigned the new species
to Pontoniopsis.

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 163-168

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

R. Michael Ewing and Daniel M. Dauer

Abstract.—A new species of Capitellidae Amastigos caperatus from the
lower Chesapeake Bay and Atlantic Ocean off the east coast of the United
States is described below. Capitomastus aciculatus Hartman 1959 is rede-
scribed. A key to the Capitellidae of the lower Chesapeake Bay is given.

Capitellidae Grube, 1862
Amastigos Piltz, 1977
Amastigos caperatus, new species
Fig. 1, a—e
Mediomastus acutus (in part).—Hartman, 1969:385-386, figs. 1-3.

Material examined.—Holotype (USNM 61107), 10 paratypes (USNM
61108), CHESAPEAKE BAY (37°07’N, 75°59'W), 6.7 m, in clean medium
sand, 19 June 1978, D. M. Dauer and staff, collectors. Ten specimens
(USNM 61109): Jones Inlet Dumpsite (40°34’N, 73°38’W), 10 m, in clean
medium sand, May 1979, Interstate Electronics Corporation, collectors for
Taxon, Inc. Other specimens: About 20 specimens from type-locality; 8
animals from Cape Henry, Virginia (36°56'’N, 75°55'W), 18.9 m, in clean fine
sand, 13 June 1979, Virginia Institute of Marine Science, collectors.

Description.—Length of holotype 18.5 mm, width 0.5 mm, 61 setigerous
segments. Lengths of 10 additional complete specimens ranged from 10.5
to 16.0 mm, widths 0.3 to 0.5 mm, setigers 34-51.

Color tan to reddish-brown in alcohol; scattered dark pigment spots along
dorsum in some specimens. Surface epithelium nearly smooth except for
Shallow transverse wrinkles of contraction; intersegmental furrows pro-
nounced in posterior setigers. Thorax and abdomen not clearly separable;
however, setigers 9 and 10 appear to be transitional in their length, number
of neurosetae per fascicle, and length of notosetae. There are no branchiae
Or Conspicuous parapodial processes; nephridial pores not observed.

Prostomium slightly depressed, triangular, tapering anteriorly to a fine tip
(Fig. la), with 2 inconspicuous nuchal slits at posteriolateral borders.
Achaetous peristomium with 2 subdermal eyespots. Eversible pharynx bul-
bous, covered with minute papillae.

164 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

|.Omm (a)

b 0.25mm (b)
[pn

Fig. 1. Amastigos caperatus: a, Anterior end in lateral view; b, Posterior end in lateral
view; c—d, Lateral and frontal view of neuropodial hooded hook from setiger 4; e, Frontal view
of notopodal hooded hook from setiger 9.

Anterior 8 setigers approximately 112-2 times wide as long; ninth setiger
distinctly longer and slightly narrower (conspicuous change in segmental
length occurring at setigers 10 or 11 in several specimens); thereafter, seg-
ments gradually increasing in length until mid-abdominal region where they
are 2—3 times as long as wide; posterior setigers becoming shorter and slight-
ly inflated in front of each segmental groove, terminating in a partly annu-
lated pygidium with cylindrical anal cirrus 2—7 times long as preceding seg-
ments (Fig. 1b). Anteriormost notopodia dorsolateral, well separated, but
approach middorsally by setiger 5-6. Neuropodia ventrolaterally throughout
body.

VOLUME 94, NUMBER 1 165

Neuropodia with 2—5 multidentate hooded hooks (Fig. Ic, d) per fasicle
through setiger 9; abrupt increase in number of neuropodial hooks to about
10 per fascicle (6—10) in tenth setiger; as many as 16 hooded hooks per
fascicle in abdominal neuropodia of several specimens, decreasing in num-
ber toward posterior but present in all segments to pygidium. These struc-
tures appear to terminate in a beak consisting of a strong, slightly curved
fang surmounted by a crest of at least 7 teeth arranged in 2 or more rows.

Notopodia with 2—3 hooded hooks per fascicle in anterior setigers, grad-
ually increasing to as many as 5 per fascicle in middle, then decreasing to
1-2 in far posterior segments. Multidentate hooded hooks of setigers 1—4
short, thick-shafted, similar in structure to neurosetae. Notopodial hooks
becoming longer and thinner through setigers 5—8; hood gradually enclosing
main fang, becoming compressed from front to back (i.e., perpendicular to
fang); margin appearing slightly crenulate. Notosetae from setiger 9 notice-
ably elongate, approximately twice as long as those of the preceding setiger
and greater than 5 times as long as notopodial hooks of setiger 1; margin of
hood distinctly wrinkled in most specimens, appearing as a bilimbate seta
with scalloped edges; aperture tightly closed around beak; fang and crest
nearly invisible in frontal view in ordinary light microscopy, appearing only
as a minute button-shaped structure at the distal end of the long and slender
shaft (Fig. le).

No sexual dimorphism was noted in a comparison of 2 ovigerous females
and a single ripe male.

Remarks.—As noted by Fauchald (1977) the exact number of thoracic
segments is often difficult to assess in the capitellids. In most specimens
examined the achaetous peristomium was not easily recognized though its
presence was confirmed through scanning electron microscopy (SEM). The
asetigerous segment (first thoracic) thus should not be regarded as a key
taxonomic character in Amastigos caperatus.

Recent detailed studies of capitellid setae (i.e., Thomassin and Picard,
1972; Warren, 1976) have shown the microstructure of hooded hooks to be
more complex than often assessed using conventional optical microscopy.
SEM techniques similar to Warren (1976) were employed in this study with
only marginal success, since the sheath surrounding the neuropodial and
anterior notopodial hooks in A. caperatus was only partly removed.

As noted earlier, the length of the main fang of notopodial hooks de-
creases in size as the shaft lengthens along posterior thoracic setigers. Our
observations with SEM indicate that the decreasing size of the beak and
hood and construction of the hood aperture around the fang is accompanied
by a reduction in the number of teeth in the crest.

Amastigos caperatus resembles the type-species, A. acutus Piltz 1977, in
that both lack capillary setae and have the characteristic flattened, modified
notopodial hooks from the posterior thoracic region. The 2 species differ in

166 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

several respects. In A. caperatus anterior setigers are relatively uniform in
width, gradually increasing in length posteriorly, with thorax and abdomen
separated by 2-3 transitional segments, whereas, in A. acutus the first 2
thoracic setigers are distinctly longer and broader than the following seg-
ments, with an abrupt transition from thorax to abdomen at setiger 9. In
addition, A. caperatus is provided with an anal cirrus while the posterior
end of A. acutus terminates in a short conical pygidium. The external ap-
pearance of the modified notosetae of A. caperatus is of considerable in-
terest; the unique wrinkled margin of the hood in most notopodial hooks is
highly consistent among individuals examined over a wide geographic range
and is an easily observed taxonomic character.

Ecology.—Amastigos caperatus occurs primarily in subtidal clean, fine
to medium sands.

Etymology.—tThe specific name refers to the wrinkled appearance of the
notopodial hoods, particularly in the abdomen.

Distribution.—Lower Chesapeake Bay; Atlantic Ocean, from Charleston,
South Carolina to Broad Sound, Massachusetts.

Capitomastus aciculatus Hartman 1959

Hartman (1959) described Capitomastus aciculatus from St. Andrews
Bay, Florida, with the setal formulae illustrated in Fig. 2 (where A = acic-
ular spine, C = capillary seta, H = hooded hook, G = genital spine, M =
mixed fascicle of capillary setae and hooks). Individuals collected from the
Chesapeake Bay did not consistently show this pattern.

Variations from the setal arrangement reported by Hartman (1959) were
as follows:

Males: 1. Some individuals had acicular spines only in the notopodia of
the first 2 setigers or lacked these spines entirely. In each case parapodia
without acicular spines had capillary setae only. There was no apparent
relationship between the setal arrangement and size although a greater pro-
portion of larger males have the acicular spines.

2. Fascicles of mixed capillary setae and hooks were occasionally found
in the neuropodia of setiger 8. |

3. Most males examined (24 of 30 specimens) lacked capillary setae in the
neuropodia of setiger 9, which according to Hartman should be mixed.

Females: 1. Individuals were also found that lacked acicular spines in the
notopodia of the first 2 setigers.

2. Several specimens had capillary setae only in the notopodia of seti-
ger 8.

When the acicular spines are absent Capitomastus aciculatus may be
confused with Capitella capitata. Males of the 2 species differ in the pres-
ence of some capillary setae in the eighth neuropodia in C. aciculatus where

VOLUME 94, NUMBER | 167

SoS a Sa ES ee ca aE
NU dalads ists ts ts [s bn} o

| CY VSM ePOS HASS T9a7/o2 KBubod

NU [stelstststs ts ts [a] ?

Fig. 2. Thoracic setal arrangement of Capitomastus aciculatus as described by Hartman
(1959).

only hooded hooks are present in C. capitata. Although Warren (1976)
noted that capillary setae rarely appear in the neuropodia of setiger 8 in C.
capitata, this situation was not encountered in 124 C. capitata specimens
examined from Chesapeake Bay. All possible setal combinations were ob-
served in the eighth neuropodia of C. aciculatus males; capillary setae may
be absent in one, but never in both, neuropodial fascicles. C. aciculatus
females have genital spines only in the notopodia of the ninth setiger and
are easily recognized since C. capitata females have only hooded hooks.

Key to the Capitellidae of the lower Chesapeake Bay*

le Camillanvasetac TeSeMt sce cara eps cars oes leah Haas so Peewee 7)
= (Cayoilleiny See BlSett Gcaocoauoscgscuscedosdes Amastigos caperatus
2. No more than 4 anterior setigers with capillary setae only ....... 3
— At least 5 anterior setigers with capillary setae only............. 5
3. First 3 setigers with capillary setae only ............ Capitella jonesi
— First 4 setigers with capillary setae only (Mediomastus) ......... 4
4. Notopodia of posterior abdominal segments with both hooks and
capillary setae or with capillary setae only ............ M. ambiseta
— Abdominal notopodia with hooks only............ M. californiensis
5. First 5 setigers with capillary setae only .... Heteromastus filiformis
— More than 5 thoracic setigers with capillary setae .............. 6
CHaGremitales HINES EPLESE Mile st boas at eter acy hace stcen lew ede ee Nc RMB Ne ex sneer eaie eas 7
SAGEM CAS PINes AWS ews Sayers 5 dusyn syt Merete as Ee SL hey 9
fea Genitalaspines presemiminseMmeensro amass os. sare aa. eee - 8

* Setal characters may vary with age (Warren, 1976). This key is based upon setal characters
of adults.

168 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

= Genital’spines present only insninth setiger =. 4-2 eee
Lh Fee Ronan eC SCM Hee Nn en Capitomastus aciculatus (female)
8. Neuropodia of setiger 8 with some capillary setae ..............
ne Meta = ete ee RD RA ge PMOL at occa tc Capitomastus aciculatus (male)
— Neuropodia of setiger 8 with hooks only ... Capitella capitata (male)
9. First 7 setigers with capillary setae or mixed capillary setae and
hoOGksSare ree ee ae oe ee ae ee Capitella capitata (female)
— First 11 setigers with capillary setae only (Notomastus) ......... 10
10. Setiger 1 with capillary setae in both noto- and neuropodia ......
Sa aV on San ui nest Biber iu eriioa bahia a cutee MSS CRD Uulbi b 0 N. latericeus
=) Mimst Scuigen withoutmeunoSsetae my e+ nce eee N. hemipodous

Acknowledgments

We would like to thank Brown and Root, Inc. of Houston, Texas for
supporting in part this research. Specimens were supplied to us by Rod
Burtelsen of the Virginia Institute of Marine Science and by Joseph Benedict
of Taxon, Inc.

The authors are especially grateful to Dr. Kristian Fauchald of the Smith-
sonian Institution for examining several of our specimens and for critically
reviewing the manuscript. We also thank Linda Lutz for preparing the fig-
ures for press.

This research was supported in part by NOAA grant NA79AA-D-00026.

Literature Cited

Fauchald, K. 1977. The polychaete worms. Definitions and keys to the orders, families, and
genera.—Nat. Hist. Mus. Los Angeles County. Science Series 28:1—190.

Grube, A. E. 1862. Noch ein Wort tiber die Capitellen and ihre Stellung im Systeme der
Anneliden.—Arch. Naturgesch. Berlin 28:366—378.

Hartman, O. 1959. Capitellidae and Nereidae (Marine Annelids) from the Gulf side of Florida,
with a review of freshwater Nereidae.—Bull. Mar. Sci. Gulf and Caribbean 9(2):153-
168.

Hartman, O. 1969. Atlas of sedentariate polychaetous annelids from California.—Allan
Hancock Found., Univ. of Southern California, Los Angeles, 812 pp.

Piltz, F. M. 1977. A new genus and species of polychaete (Family Capitellidae) from southern
California.—Bull. So. Calif. Acad. Sci. 76(1):57—60.

Thomassin, B., and C. Picard. 1972. Etude de la microstructure des soies de polychétes
Capitellidae et Owentidae au microscope électronique a balayage: un critere systema-
tique précis.—Mar. Biol. 12:229-236.

Warren, L. M. 1976. A review of the genus Capitella (Polychaeta: Capitellidae).—J. Zool.,
Lond. 180:195—209.

Department of Biological Sciences, Old Dominion University, Norfolk,
Virginia 23508. RME present address: Barry A. Vittor & Associates, Inc.,
8100 Cottage Hill Road, Mobile, Alabama 36609.

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 169-180

UCA MARGUERITA, A NEW SPECIES OF FIDDLER
CRAB (BRACHYURA: OCYPODIDAE) FROM
EASTERN MEXICO!

Carl L. Thurman II

Abstract.—A species of fiddler crab new to science, Uca marguerita, on
the eastern terrigenous coastline of Mexico is reported. Previously, several
specimens of the newly described species were confused with either Uca
virens Salmon and Atsaides, Uca burgersi Holthuis, or Uca speciosa Ives.
A detailed morphological description of adult specimens along with a general
description of the male’s courtship display is presented. The new species
is compared with sympatric congeners. On both morphological and behav-
ioral grounds, Uca marguerita appears to be a member of the subgenus
Minuca of Crane (1975).

Several authors have reviewed the North American fiddler crabs. How-
ever, only Rathbun (1918), Buitendyk (1950), and Crane (1975) have re-
ported new records from the east coast of Mexico. Rathbun identified Uca
mordax Smith in collections from Tampico, Mexico. Buitendyk also be-
lieved U. mordax as well as U. speciosa Ives were present in the Mexican
state of Veracruz. Recently, Holthuis (1967) and von Hagen (1970) consid-
ered Buitendiyk’s U. mordax records to be Uca burgersi Holthuis. In a
major taxonomic revision, Crane reclassified Rathbun’s Mexican specimens
as either Uca vocator Herbst, Uca spinicarpa Rathbun or Uca pugnax
virens Salmon and Atsaides. Simultaneously, she also added 20 new spec-
imens which she believed to represent U. p. virens from southern Veracruz.
These gifts to the New York Zoological Society extended the presumed
range of Uca virens from the area between Ocean Springs, Mississippi, and
Port Aransas, Texas (Salmon and Atsaides, 1968), into southeast Mexico.
Uca rapax Smith was also recognized to range as far north as Tamaulipas.

The taxonomic assessments of both Crane (1975) and Holthuis (1967) are
based on examination of a few preserved museum specimens from eastern
Mexico. In addition several authors have expressed the opinion that the
original type characteristics for Uca virens as well as Uca longisignalis
Salmon and Atsaides are unreliable (Crane, 1975:203; von Hagen, 1975,
1980). On the one hand, Crane tentatively proposed five traits to distinguish

! Adapted as part of a doctoral dissertation submitted to the Graduate School, University of
Minnesota, Minneapolis.

170 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

U. virens from other Minuca but felt her choice to be marginally acceptable
without a more complete collection series. On the other, von Hagen’s (1980)
approach has been to synonymize the newer species with older, traditionally
described taxa. Complete resolution of this controversy is beyond the scope
of the present study. However, as a result of field collections and exami-
nation of available preserved museum specimens, it is apparent that Crane’s
difficulty in obtaining reliable traits for U. virens arises from the simulta-
neous treatment of at least two species. Part of the U. pugnax virens as
well as some of the U. burgersi and U. speciosa from eastern Mexico are
members of an undescribed species. This new species, Uca marguerita, is
limited to the terrigenous areas of eastern Mexico. It does not occur in the
United States or upon the biogenic soils of the Yucatan peninsula.

Materials and Methods

Populations of fiddler crabs in the western and southern Gulf of Mexico
were studied during December 1972, September 1974, March 1976 and Sep-
tember 1976. More than 3000 specimens were collected from 77 localities
between Ocean Springs, Mississippi, and Chetumal, Quintana Roo, Mexico.
Two hundred thirteen individuals of the new species were found in localities
between the states of Tamaulipas and Campeche, Mexico. From these col-
lections, lots were deposited in the USNM, Department of Invertebrate
Zoology, Washington, D.C. Samples of the new species were supplied from
Tamaulipas, La Pesca; Veracruz, Rio Panaco; Veracruz, La Barra; Vera-
cruz, Nautla; Veracruz, Laguna La Mancha; Veracruz, Boca del Rio; Ta-
basco, Puerto Ceiba; Tabasco, Frontera; Campeche, Rio San Pedro. The
specimens from Laguna La Mancha in Veracruz are designated as male
holotype (USNM 180400) and female allotype (USNM 180401). The re-
mainder (USNM 180443 through 180450) are designated as paratypes. In
addition, older preserved materials also were examined. The following Min-
uca were made available by the USNM:

Uca burgersi Holthuis 1967—138490 (Belize), 139173 (Belize), 13488
(Florida).

Uca longisignalis Salmon and Atsaides 1968—121599 (Holotype) 122204
(Paratype), 2259 (Grand Isle), 21845 (Biloxi), 33035 (Matagorda), 72132 part
(Texas), 74902 (Ocean Springs), 122764 (Yankeetown), 138647 (Yankee-
town), 138648 (Galveston).

Uca minax (LeConte 1855)—42618 (Sarasota), 64083 (Ft. St. Phillip),
64155 (Grand Isle), 98144 (Lake Pontchartrain).

Uca mordax (Smith 1870)—18430 (Swan Island), 50950 (Belize).

Uca pugnax (Smith 1870)—17488 (Winyah Bay), 71196-7 (Ft. Macon),
74455 (Brigantine).

VOLUME 94, NUMBER 1 171

Uca rapax (Smith 1870)—15057 (Mobile Bay), 25033 (Matagorda), 95526
(Yucatan), 96475 (Laguna Madre).

Uca virens Salmon and Atsaides 1968—121598 (Holotype), 122205 (Para-
type), 30570 (Cameron), 33031 (Matagorda), 72132 (Texas), 122765 (Port
Aransas), 72131 (Texas), 138642 (Corpus Christi), 138643 (Port Aransas),
43353 (Tampico), 138644 (Veracruz), 138645 (Veracruz), 18689-dry (Tam-
pico).

Uca vocator (Herbst 1804)—21373 (Belize), 138855 (Veracruz), 139175
(Tampico).

The following lots were made available by the Rijksmuseum:

Uca burgersi Holthuis 1967—7577 (Veracruz), 7578 (Veracruz).

Uca speciosa (Ives 1891)—7574 (Veracruz).

A number of morphological characteristics were measured in order to
characterize the new species. Measurements were taken to the nearest 0.05
mm with a vernier caliper. Carapace breadth was measured at maximal
distance between anterior dorsolateral margins. Body length was taken from
the front of carapace to posterior margin above abdomen. Frontal width
was taken between the inner junctions of the eyebrows with the lower fron-
tal margin of the carapace. The length and width of the merus was measured
along the respective central axis of the fourth ambulatory segment. Propodal
ratios were obtained from measurements taken on the fifth segment of the
ambulatory. The details of gonopodium and gonopore structures were stud-
ied with the aid of a stereo dissecting microscope.

The cheliped waving behavior of male fiddler crabs was recorded in the
field using a 16 mm movie camera where ambient temperatures were always
above 25°C. General wave form and temporal components of the courtship
display were determined by frame analysis. Since this behavior was record-
ed at either 16 or 24 frames per second, estimates of temporal characteristics
were determined by multiplying the number of frames involved by either
0.0625 or 0.0416 seconds for respective recording speeds.

Results
Uca marguerita, new species
Figs. 1, 2

Uca mordax.—Rathbun, 1918:391—393 [part].

Uca mordax.—Buitendik, 1950:279 [part].

Uca speciosa.—Buitendik, 1950:279 [part].

Uca burgersi.—Holthuis, 1967:51—54 [part].

Uca burgersi.—von Hagen, 1970:225 [part].

Uca speciosa.—von Hagen, 1970:227 [part].

Uca pugnax virens.—Crane, 1975:200—205 [part].—not Uca virens Salmon
and Atsaides, 1968.

172 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Uca marguerita: A, Dorsal view; B, Inner surface of male cheliped; C, Ventral
view; D, Frontal view of interocular region. White bar = 1 cm.

Type-locality.—Mexico, Veracruz, Laguna La Mancha (19°40'’N, 96°20’ W)
between Villa Rica and Cardel. In addition to the USNM paratypes, ex-
amples from the following lots have been identified as Uca marguerita:
USNM 43353 part, USNM 138644, USNM 138645, RMNH 7574 part,
RMNH 7577 part, and RMNH 7578 part.

Diagnosis.—Fig. 1. Width of rounded frontal lobe less than 4% maximum
carapace width. Ventral margins of distal ambulatory segments without pu-
bescence. In males, superior carina above carpal cavity in palm of cheliped
consists of a row of tubercles. Distal end of this carina turned forward
toward dactyl. In females, genital operculum without prominent tubercle.

Etymology.—Marguerita a common christian name used in the endemic
region of the crab.

Morphological description.—From dorsal view (Fig: 1A), carapace length
66.0% or less of breadth. Carapace widest slightly posterior to anterior
margins. Largest male 24.7 x 15.1 mm; largest female 17.3 x 12.0 mm at
Nautla, Veracruz. Surface finely granular and pitted in both sexes. Females
have few large tubercles in epibranchial regions. Neither pubescence nor
numerous setae on dorsal carapace. H-depression in cardiac-mesogastric
region very prominent. Color in preserved specimens homogeneously dark
except for occasional concentration in depression. Carapace of living spec-
imens brown, dark green, or gray. Frontal lobe between eyestalks less than
33.0% of maximum carapace width (Fig. 1D); in males near 27%, in females
near 31%. From dorsal view, frontal lobe distinctly rounded or spatulate

VOLUME 94, NUMBER I 173

and very shallow. Maximum width of eyebrow almost equal to diameter of
adjacent eyestalk peduncle. Distal cornea occupies between 25 and 33% of
peduncle length. Anterior margins finely granular in both sexes. Sulcus pos-
terior to eyebrow forms deep transverse depression curving toward meso-
gastric region. Anteriorly, dorsolateral margins of carapace curving inward
slightly. Junction of anterior and posterior forming curved but obtuse angle.
Posteriorly lateral margins converging toward midline of carapace giving
body shortened appearance.

When carapace is viewed from anterior (Fig. 1D), lobed interocular region
very shallow and rounded. Carapace moderately arched through axis of
width. From lateral view, carapace strongly convex, almost subcylindrical.
Eyebrows strongly inclined horizontally.

Outer and upper manus of cheliped covered with very large tubercles.
Dorsal margin distinctly tuberculate. Ventral margin of manus with tuber-
cles forming a distinct keel. Submanus sulcus prominent. Tubercles of outer
manus becoming minute on pollex. Both pollex and dactyl flattened, more
bladelike than tubular (Fig. 1B). Pollex with 3 rows of tubercles in gape.
Usually 1 large tubercle or ‘‘tooth’’ more than halfway along pollex in gape.
Three to 5 tubercles at distal end of pollex. Tip thick. Pollex with 4 or 5
large tubercles on ventral margin of dactyl-propodus articulation that do not
traverse from outer to inner pollex surfaces. Dactyl with teeth in gape.
Upper margin of carpal cavity on inner manus lined with a prominent ridge
of tubercles or superior carina (Fig. 1B). Some pubescence may be present
in cavity. Distal end of carina turned distinctly toward dactyl as it ap-
proaches palm. Region dorsal to end of carina forming a smooth, triangular
sulcus. Ventral manus with oblique tuberculated ridge extending about %4
distance to pollex. Distally, this consists of 2 or 3 ill-defined rows of smaller
tubercles. Proximately, these converge into a prominent, elevated apex con-
sisting of larger tubercles at ventral border of carpal cavity. Apex vertical
to surface of palm. From apex, a row of tubercles follows outer edge of
carpal cavity and extends upwards, almost intersecting with superior carina.
Proximal palm covered with numerous large tubercles equal in size to those
on oblique ridge. Palm area appears swollen because of deep sulcus at in-
tersection of pollex and high apex of oblique ridge. Two rows of tubercles
near dactyl articulation: one row on predactyl ridge, other more proximal
to palm. Proximal row divergent from distal with dorsal end curving toward
dactyl.

Form of minor cheliped similar to that of other Minuca. Distal end of
pollex and dactyl sparsely setose. Usually fewer than 10 short hairs on
either finger. Serrations in gape weakly developed. Distal end of merus with
a few long setae parallel to dorsal propodal articulation.

Dorsal margins of ambulatory merus slightly convex. Both anterior and
posterior surfaces covered with short setae. Merus of third ambulatory

174 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

[0.5 mm ———|

RAPAX MARGUERITA BURGERSI

Fig. 2. Gonopodium structures of Uca rapax Smith, Uca burgersi Holthuis and Uca mar-
guerita from eastern Mexico. Anterior view of right gonopodium from males of carapace widths
17 to 18 mm. 60x.

about 2.56 times longer than wide in males and about 2.22 times in females.
Ventral surface of first, second, and third meri with numerous setae but no
pubescence (Fig. IC). Usually at least 5 setae per merus. Fourth leg with
0 to 5 setae per merus. Propodal and carpal segments of first 3 ambulatories
with pubescence only on dorsal surfaces. Occasionally pubescence present
on fourth ambulatory propodus and carpus, more frequent in females than
males. Numerous long setae present in rows on anterior, posterior, dorsal
and ventral surfaces of propodus. Two rows on dorsal surface. Propodus
width about 30% of length on first ambulatory in males.

Copulatory pleopods of male thin and nearly straight when viewed from
sternum. After terminal setae are removed, distal end appears constricted
before angling of tip (Fig. 2). Tip turned almost to right angle with lateral
margins of carapace. Distal tip with small shield-shaped cuticular process.
Free edge smooth and undeveloped. Two soft palps present. One subter-
minal near constriction extending distally to near origin of cuticular process.
Second palp terminal on inner lateral surface with distal end embracing
cuticular process. Figure 2 compares tip of nude gonopodium of Uca mar-
guerita with those of U. burgersi and U. rapax of comparable size. These
differences appear consistently across the range of the new species. Female
gonopore without tubercle. Operculum lined with thin rim from sternum.
Anterior margin may show small microscopic fold with edge protruding
toward center of operculum.

Geographical distribution.—The new species was collected in brackish

VOLUME 94, NUMBER 1 175

SIE GOINIDS

Fig. 3. High intensity courtship display in male Uca marguerita. Major positions of che-
liped indicated by parentheses. (See text.)

habitats between La Pesca, Tamaulipas, and Rio San Pedro, Campeche,
Mexico. The specimens reported as Uca pugnax virens from Mexico
(Crane, 1975; USNM 43343 part, 138644 and 138645) belong to the new
species. The additional 16 specimens from Rio Tonchochapa could not be
located. Holthuis (1967) and von Hagen (1970) both consider the entire lots
RMNH 7577 and 7578 to be U. burgersi from Veracruz. One male from
RMNH 7577 (Nautla) and 3 males and 1 female from RMNH 7578 (Casitas)
are Uca marguerita. The clawless male from U. speciosa lot RMNH 7574
is also the new species (Casitas). The new species thus appears to be en-
demic to the western shores of the Bay of Campeche.

Male courtship display.—The cheliped waving of the new species has
been recorded at two localities within its geographic range. Temporal and
topological components of this supplementary characteristic are useful for
distinguishing the new species in the field and aiding in the determination
of phylogenetic affinities. Thirty waves from seven males at La Pesca, Ta-
maulipas, and Puerto Ceiba, Tabasco, were studied by frame analysis.

The form of the wave in the new species is relatively simple and appears
consistent between northern (n = 2) and southern (n = 5) males. As illus-
trated in Fig. 3, the heel of the cheliped (proximal manus) and carpus are
raised to a level near the front of the carapace (position 1 and 2). The fingers
of the claw then follow and are held momentarily at the carapace front
(position 2). At this point, the axis of the manus and dactyl are more or less
parallel with the substrate. The cheliped is oscillated laterally while in this
position (3). The propodus is then quickly flexed vertically through a vertex
(position 4) and downward (position 5) to a frontal position (6). There is no
apical pause but the cheliped is rotated slightly exposing the inner manus
during the apical transition. On the rapid return or downstroke, the fingers
of the claw do not touch the substrate. Also during the descent phase, the
proximal carpal segment is lowered before the distal manus and dactyl (po-
sition 5).

The singular frontal pause appears to be characteristic of the wave. Only

176

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Temporal and topological characteristics of waving behavior in Uca marguerita
from Eastern Mexico.

Mean

Type duration hold Pause To apex Return
of wave No. (sec) SE (sec) SE (sec) SE (sec) SE (sec) SE
High 6 0.92 +0.14 0.41+0.05 0.14+0.05 0.12 + 0.08 £0.28 + 0.10
Moderate 19 1.54 = 0.21 O76%== 012255 0:19 ==70:309 © 023 = 0109 0 SS 0sI4
Low 5 2.63 + 0.61 1366) 2210257 79 70332 == 013) 1018) 2009) FOSS OaS
0.18 + 0.06 0.34 + 0.06
Duration between
waves (sec)
High 0.80 + 0.06 (n = 4)

Moderate
Low

1835 p=1 O22 (nal)
3.28 + 1.40 (n = 5)

one individual waving at low intensity expressed two pauses. Pauses at the
apex or on the return flexure were never noted. During the upward move-
ment of the claw after the frontal pause, the second and third ambulatories
on the minor side and the second on the major may be extended laterally
to a position parallel with the substrate. These are returned to supporting
positions before the cheliped is brought down to the frontal position (6).
The minor cheliped is also flexed laterally after the initial pause and oscil-
lation of the large cheliped.

The wave displays may be classified into three intensities on the basis of
duration (Table 1). The highest intensity waves are observed with the ap-
proach of a female. These waves are approximately 1.0 s in duration. Low
intensity waves are a little more than 2.5 s. The latter were observed when
no other crabs were near. The time course of successive waves changes
with the approach of another crab. At first, waves are expressed very rapidly
and the upward pausing phase (position 2 and 3) is scarcely discernable. As
a female approaches, durations are then increased and a bobbing or curtsy
(Crane, 1975:496, 658) is incorporated. During this slower waving, the legs
may or may not be extended laterally. A

The movement of the cheliped from an initial pause to a frontal position
at the completion of a wave does not change with increase in waving inten-
sity (Table 1). The complete execution from initial pause to unflexed frontal
position occurs in approximately 0.5 s without stop at vertex. Apparently
wave duration is determined by the amount of time spent raising the heel
and dactyl to a frontal position (1 & 2) and time spent in oscillation pause
phase (3). The initial step may vary from 1.66 to 0.41 s while the pause may
simultaneously vary from 0.32 to 0.17 s between low and high intensity
waving.

VOLUME 94, NUMBER 1 177

On the other hand, pauses between successive waves are proportional to
wave duration. During high intensity waving, interwave duration is 0.80 s.
At intermediate intensity, the interval is increased to 1.35 s while at low
intensity there are 3.28 s between waves. The lowest intensity waves are
extremely variable in this component.

Discussions and Conclusions

The general morphology of the new species is most closely allied with the
fiddler crab subgenus Minuca (Crane, 1975). In the Atlantic, this taxon
possesses two more or less related superspecies, galapagensis and minax,
as well as the species U. burgersi. Of the superspecies, galapagensis pos-
sesses frontal breadths less than 14 carapace width while the minax super-
species and U. burgersi have frontal-carapace relationships exceeding this
value. Consequently, Uca marguerita may be distinguished from the broad-
fronted western Atlantic Uca mordax, U. pugnax, U. longisignalis, U. vo-
cator, U. burgersi, and U. minax. Moreover, it can be separated from U.
mordax and U. longisignalis by a lack of pubescence on the ventral surfaces
of ambulatory meral, carpal and propodal segments (Crane, 1975:630-—631).
Females also lack a genital tubercle, which aids in discerning them from U.
mordax, U. pugnax, U. vocator, U. minax and U. longisignalis as well as
the narrow-fronted U. rapax.

The Uca pugnax virens of Crane from Mexico (USNM 138645) do not
possess the genital tubercle. On the other hand, the only female specimen
claimed by Salmon and Atsaides (1968) as a U. virens (USNM 72132) has
this structure. Also, the female from Crane’s Texas lot (USNM 72131) which
is believed to be U. p. virens is equipped with the genital tubercle. The
Mexican samples of females discussed by Crane are thus quite distinct from
the Texas samples of U. virens, and are considered to be Uca marguerita.
By the same token, the largest female of the U. burgersi sample (Holthuis,
1967) from Mexico, in which all lack genital tubercles, possesses a frontal
width less than 4 carapace width. This is also U. marguerita.

Of the Atlantic Minuca, U. marguerita was found sympatric with only
U. burgersi, U. vocator and U. rapax. The males of the new species are
readily distinguished from U. vocator on the basis of absence of pubescence
from the carapace and dorsal surfaces of ambulatory meri, a highly devel-
oped oblique ridge across the palm of the major cheliped, and a short ap-
pearing body. The new species is more difficult to distinguish from the
galapagensis superspecies and U. burgersi. However, it may be segregated
from U. burgersi by its narrower frontal region, and a more granular, shorter
appearing carapace. On the inner surface of the male’s large cheliped (Fig.
1B), the tubercles of she superior carina along the upper margin of the
cheliped carpal sinus turn toward the dactyl in U. marguerita, but back to

178 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

the edge of the cavity in Uca burgersi. On the inner palm, the proximal
predactyl ridge is not as arched in U. marguerita as in either U. burgersi
or U. mordax. Although U. burgersi and U. marguerita possess similar
gonopores, the male gonopods are distinct in their areas of sympatry (Fig.
2). Those in the new species are thinner, more straightened proximately and
have a more sharply angled tip than U. burgersi. The cuticular tip is shorter
in the new species while the proximal and medial palps appear longer in U.
burgersi.

Unfortunately, the general dimensions of the carapace and frontal lobe in
males of both Uca rapax and the new species are quite similar. The frontal
lobe of the new species is distinctly spatulate as opposed to the angular or
truncated lobe of U. rapax. The eyebrows are broader in Uca marguerita
than in either U. rapax or U. pugnax. In U. marguerita the eyebrows are
horizontally inclined to such a degree that they appear almost as wide as
the adjacent eyestalk peduncle. The eyebrows of U. rapax as well as U.
virens (Crane, 1975:201) are sufficiently angled so that they are clearly vis-
ible in a dorsal carapace view. Since the eyebrows of U. pugnax are nearly
vertical in a lateral view they are barely visible from the median dorsal
perspective (Rathbun, 1918:395, 398). The cheliped contains tubercles that
are large and an arched, proximal predactyl ridge that is distinct from the
smooth palm and parallel ridge of male U. rapax or U. virens. The oblique
ridge of U. marguerita is developed to a much higher apex than in other
local Minuca. The distal extension of the superior carina is also distinct
from those found in both U. rapax and U. pugnax. The gonopod of U.
rapax is distinctly broader and possesses a more extended cuticular tip than
in the new species (Fig. 2). The male gonopod was helpful in identifying the
clawless U. marguerita in RMNH 7574.

The waving display of the new species also is distinct from other Atlantic
Minuca. These are similar to those expressed by other members of the
galapagensis superspecies. The duration of single waves in U. longisig-
nalis, U. minax, U. pugnax, U. vocator and U. burgersi are considerably
longer than the waves of the new species. In particular, the duration of the
lowest intensity waves in the new species is approximately equal to the high
intensity waves of both Uca rapax (Salmon, 1967; Crane, 1975) and U.
virens (Salmon and Atsaides, 1968). Moreover, the ascending portion of
waves contains fewer ascending pauses than any other Atlantic Minuca. On
the basis of ethology, the new species is obviously readily distinguishable
in the field.

Relationships.—Although less is known about the Pacific Minuca, von
Hagen (1968) described in some detail the waving behavior and gonopod
structure of both U. galapagensis Rathbun and U. herradurensis Bott from
Peru. These Pacific relatives differ from U. rapax and U. pugnax by having
the palm of the cheliped tuberculate, the oblique ridge of the claw developed

VOLUME 94, NUMBER | 179

to a high apex and strong postorbital sulcus on dorsal carapace. Both are
moderately narrow-fronted Minuca (Crane, 1975:183). The genital tubercle
is absent in the female of the Pacific species. The same ensemble of char-
acters may be used to distinguish the new species from its Atlantic relations.

Although the waves of male Uca galapagensis are highly variable, they
are distinct from those of U. rapax and U. herradurensis. In U. galapa-
gensis, vibrational components accompany a single apical pause. Although
the new species lacks any pause at the vertex, the positioning of the cheliped
in front of the mouth parts at the end of a wave, oscillatory movements,
and short wave durations in U. marguerita courtship display are reminiscent
of the patterns in Uca galapagensis (von Hagen, 1968; Crane, 1975). Con-
sequently, on the basis of similarity in morphology and behavior, Uca mar-
guerita, the new species, tentatively appears to be phyletically close to the
Pacific relatives in the Minuca superspecies, galapagensis.

Acknowledgments

I wish to thank Dr. F. H. Barnwell, of the University of Minnesota, for
the use of many preserved specimens, and his comments and support of
this research. I also would like to thank Drs. Raymond B. Manning, U.S.
National Museum of Natural History, Smithsonian Institution (USNM), and
L. B. Holthuis, Rijksmuseum van Natuurlijke Historie at Leiden (RMNH),
for making preserved specimens available. Financial support for field studies
was provided by the Dayton Natural History Fund from the Bell Museum
of Natural History, University of Minnesota.

Literature Cited

Buitendijk, A. M. 1950. Note ona collection of decapod Brachyura from the coasts of Mexico,
including the description of a new genus and species.—Zool. Meded. (Leiden) 30:269-
282.

Crane, J. 1975. Fiddler Crabs of the World (Ocypodidae; genus Uca).—Princeton University
Press, Princeton. 1-736.

von Hagen, H. O. 1968. Studien an peruanischen Winkerkrabben (Uca).—Zool. Jahrb. Sys-

tem. 95:395-468.

1970. Verwandtschaftliche Gruppierung und Verbreitung der karibischen Winker-

krabben (Ocypodidae, Gattung Uca).—Zool. Meded. (Leiden) 44:217—235.

. 1975. Klassifikation und phylogenetische Einordnung der Lautausserungen von Ocy-

podiden und Grapsiden (Crustacea, Brachyura).—Z. f. Zool. System. Evol. 13:300-316.

. 1980. A key to the **x-species’’ of North American fiddler crabs (genus Uca).—Zool.

Meded. (Leiden) 55:87—96.

Holthuis, L. B. 1967. On a new species of Uca from the West Indian region (Crustacea,
Brachyura, Ocypodidae).—Zool. Meded. (Leiden) 42:51—54.

Rathbun, M. J. 1918. The grapsoid crabs of America.—Bull. U.S. Natn. Mus. 97:1—461.

Salmon, M. 1967. Coastal distribution, display and sound production by Florida fiddler crabs
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, and S. P. Atsaides. 1968. Behavioral, morphological and ecological evidence for two
new species of fiddler crabs (genus Uca) from the gulf coast of the United States.—
Proc. Biol. Soc. Wash. 81:275—290.

Zoology Program, Department of Ecology and Behavioral Biology, Uni-
versity of Minnesota, Minneapolis, Minnesota 55455. Present address: De-
partment of Medicine, The Medical College of Ohio, Toledo, Ohio 43699.

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 181-227

ADDITIONAL ACANTHONOTOZOMATID,
PARAMPHITHOID, AND STEGOCEPHALID
AMPHIPODA FROM THE SOUTHERN OCEAN

Les Watling and Heather Holman

Abstract.—Odius antarcticus n. sp. is described and 2 new subspecies of
Gnathiphimedia sexdentata are named. In addition 14 acanthonotozomatid,
5 paramphithoid, and 4 stegocephalid species are recorded, with most re-
described. [phimediella discoveryi Watling and Holman is herein considered
Synonymous with Guathiphimedia macrops. A combined key to Guathi-
phimedia and Iphimediella is given. In the genus Epimeria, E. excisipes is
placed into synonymy with E. georgiana and Subepimeria geodesiae with
FE. puncticulata. The boreal North Pacific Uschakoviella echinophora is
recorded from Antarctic waters for the first time.

This paper is the second in a series directed toward the redescription of
poorly known or incorrectly described species as well as the description of
new species. In the first paper (Watling and Holman, 1980) we described 6
new species and one new genus of Acanthonotozomatidae, 2 new species
of Paramphithoidae and one new species of Stegocephalidae, chiefly from
the Scotia Sea region. New revisions were also offered for the acanthono-
tozomatid genera Acanthonotozomella, Iphimedia, Iphimediella and Pseu-
diphimediella, and the paramphithoid genera Epimeria and Parepimeria.
The present paper contains one new species (Odius antarcticus), two new
subspecies of Gnathiphimedia sexdentata, and redescriptions of 10 acan-
thonotozomatids, 5 paramphithoids, and 2 stegocephalids. Several other
species are also recorded.

Acanthonotozomatidae
Acanthonotozomoides Schellenberg

Acanthonotozomoides Schellenberg, 1931:124.

Type-species.—A. sublitoralis Schellenberg, 1931 (original designation).

Diagnosis. —Mandible with narrow, slightly toothed apex; maxilla | palp
biarticulate, inner plate minute; maxilliped palp exceeding outer plate, 4-ar-
ticulate, article 2 slightly produced medially; gnathopods simple; coxa 1
distally acute.

182 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Acanthonotozomoides oatesi (K. H. Barnard 1930)

Acanthonotozomella oatesi K. H. Barnard, 1930:346, figs. 20, 21.—K. H.
Barnard, 1932:117, fig. 65.

Acanthonotozomoides oatesi.—J. L. Barnard, 1969:119.—Bellan-Santini,
1972:167, pl. 1.

Material.—Eltanin Cruise 12, Sta. 1003, 15 Mar. 1964, 62°41'S, 54°43’W,
210-220 m, | juvenile.

Diagnosis.—Pereonite 1 with single pair of sub-dorsal processes; coxa |
not bidentate distally.

Distribution.—From Adelie Coast, along western Antarctica to Palmer
Archipelago and South Georgia, 82—236 m.

Anchiphimedia K. H. Barnard
Anchiphimedia K. H. Barnard, 1930:357.

Type-species.—A. dorsalis K. H. Barnard, 1930 (original designation).

Diagnosis.—Upper lip incised; mandibles meet along frontal plane with
cutting edge angled downwards and toward posterior of body; maxilla 1 palp
biarticulate, very short, outer plate bears numerous curved setae instead of
strongly chitinized, toothed spines; maxilliped palp article 4 very small,
partly covered by hood-like extension of article 3, article 2 medially ex-
panded but not produced along article 3; gnathopods | and 2 chelate.

Anchiphimedia dorsalis K. H. Barnard 1930
Figs. 1, 2

Anchiphimedia dorsalis K. H. Barnard, 1930:357, figs. 29, 30.—K. H. Bar-
nard, 1932:123.

Material.—Eltanin Cruise 27, Sta. 1924, 27 Jan. 1967, 75°11'S, 176°13'W,
728-732 m, 1c.

Diagnosis.—As for genus.

Description.—The following supplements the description of K. H. Bar-
nard (1930). Upper lip incised. Right mandible with prominent, toothed ac-
cessory plate; left mandible with minute accessory plate. Maxilla 1 outer
plate slender, with numerous non-plumose setae distally; inner plate sub-
acute apically, with approximately 15 plumose setae on distal half of medial
margin. Maxilla 2 inner plate wider than outer, both densely setose distally,
setae on outer plate slightly longer than those on inner plate; inner plate
with plumose setae along medial margin. Maxilliped outer plate broad; palp
article 4 minute, articles | and 3 subequal, article 2 expanded medially and
only slightly shorter than either articles 1 or 3. Gnathopod 1 chelate; lateral
margins of articles 5 and 6 setose; article 3 one-third length of article 6.

VOLUME 94, NUMBER | 183

704)

Yip pe
Lid

Fig. 1. Anchiphimedia dorsalis, Eltanin Sta. 1924: a, Body side view; b, Antenna | pedun-
cle; c, Right mandible; d, Maxilla 1; e, Maxilla 2; f, Maxilliped.

Gnathopod 2 chelate; lateral margins of articles 5 and 6 more setose than
on gnathopod 1; articles 5 and 6 subequal, longer than article 3.

Distribution.—Palmer Archipelago along western Antarctic to McMurdo
Sound, 259-732 m.

184 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Anchiphimedia dorsalis, Eltanin Sta. 1924: a, Gnathopod 1, b, Gnathopod 1 pro-

podus and dactylus; c, Gnathopod 2; d, Gnathopod 2 propodus and dactylus without setae;
e, Telson.

Remarks.—The orientation of the mandible in this species is intermediate
between the condition seen in species of Iphimediella and that seen in Pseu-
diphimediella (documented by Watling and Holman, 1980). The cutting
edges of the incisors meet in a plane oriented more to the frontal than to
the transverse plane.

Echiniphimedia K. H. Barnard
Echiniphimedia K. H. Barnard, 1930:358.

Type-species.—Iphimedia hodgsoni Walker, 1906.

Diagnosis (from description in J. L. Barnard, 1967).—2 or more coxal
plates with submarginal fixed teeth; some or all body segments covered with
rows or groups of erect cusps on lateral surfaces; epistome broad; maxilla
1 palp biarticulate; maxilliped palp 4-articulate, article 4 extremely minute;
gnathopods 1 and 2 chelate.

Echiniphimedia echinata (Walker 1906)

Iphimedia echinata Walker, 1906:150; 1907:28, pl. 10, fig. 16.—Chevreux,
1912:119.

VOLUME 94, NUMBER I 185

Echiniphimedia nodosa.—kK. H. Barnard, 1930:361, fig. 33.

Echiniphimedia echinata.—K. H. Barnard, 1932:126.—Nicholls, 1938:80,
fig. 42.—J. L. Barnard, 1967:9, figs. 4, 5.

Pariphimediella echinata.—J. L. Barnard, 1964:51.

Material.—Eltanin Cruise 32, Sta. 1995, 10 Jan. 1968, 72°04’S, 172°38’E,
360-342 m, | juvenile; Cruise 32, Sta. 2065, 26 Jan. 1968, 78°23'S, 173°06' W,
473-475 m, | juvenile.

Diagnosis (from description in J. L. Barnard, 1967).—Pereonites 5-7 (oc-
casionally 2—7), pleonites 1-4, coxae 5-7 (occasionally 3-7), and bases of
pereopods 5-7 sparsely covered with small, fixed, marginal and submarginal
teeth; pereonite 1 smooth; antenna | peduncle article 1 lacking strong distal
teeth; maxilla 2 outer plate lateral margin with only 1 or 2 setae.

Distribution.—Davis Sea, Adelie Coast, Ross Sea and western Antarctica
to Palmer Archipelago, South Orkneys and South Georgia, 10—585 m.

Remarks.—The specimen from Sta. 2065 (8 mm) possessed teeth begin-
ning on pereonite 2 hind margin, similar to specimens reported by Nicholls
(1938).

Echiniphimedia hodgsoni (Walker 1906)

Iphimedia hodgsoni Walker, 1906:152; 1907:30, pl. 11, fig. 18.
Echiniphimedia hodgsoni.—K. H. Barnard, 1930:359, fig. 31.—Nicholls,
1938:82, figs. 43-44.—J. L. Barnard, 1967:3, figs. 1-3.

Material.—Eltanin Cruise 12, Sta. 1003, 15 Mar. 1964, 62°41'S, 54°43’W,
210-220 m, | juvenile.

Diagnosis (from description in J. L. Barnard, 1967).—Pereonites 1-4
densely covered with elongate teeth; pereonites 2—6 with 2 vertical rows of
very slender teeth; coxa 3 with more than S teeth.

Distribution.—From Davis Sea, around the western Antarctic coasts to
South Orkney Islands and South Georgia, 20-1120 m.

Gnathiphimedia K. H. Barnard
Gnathiphimedia K. H. Barnard, 1930.

Type-species.—G. mandibularis K. H. Barnard, 1930.

Diagnosis (modified from J. L. Barnard, 1969).—Antenna | with uniar-
ticulate accessory flagellum; upper lip entire or sinuous; epistome broad-
ened, greater than 3 times as broad as high; mandible short, incisor thick-
ened, oriented to cut in transverse plane; lower lip without inner lobes,
distal apices broadly rounded; maxilla 1 palp biarticulate, reaching end of
outer plate; maxilliped palp 4-articulate, article 2 slightly expanded, not
produced along article 3, article 4 minute, covered by hood-like extension
of article 3; gnathopods | and 2 chelate; telson slightly cleft.

186 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Remarks.—This genus and Iphimediella are externally very similar but
differ in the width of the epistome and shape of the mandible, suggesting
two quite different modes of feeding. The incisor of [phimediella is much
thinner indicating a cutting or tearing function whereas the incisor of Gna-
thiphimedia 1s thickened, indicating a crushing function.

Key to species of Gnathiphimedia and Iphimediella

1. Pleonites with enlarged mid-dorsal carinae, paired dorsal processes

NOtTPrOMINENE sierrphscsess ae ee I. imparidentata (Bellan-Santini)
— Pleonites with distinct paired dorsal processes ................. 2
2. Pleonites without paired dorsal processes on pleonite 3.......... 3
— Pleonites with paired dorsal processes on pleonite 3 ............ 7

3. Pereonites 6 and 7 with paired dorsal processes ................
ee ee Le ae OTR I. octodentata (Nicholls)

— Pereonite 6 without paired dorsal processes .............4.:5-5 4
4. Coxa 1 rectangular, smoothly rounded anterodistally; upper lip

TTMCISC Ch 5 Ges, 5 Fe ase ens cea aie days Gi RO Shee Geen ee acl bee 5
=) Coxa itapenne distally supper lip entine= s- 4.4 6
5. Pleonite 3 with slightly projecting keel, dorsal processes meet to

1) Gia. War? aed enor CMe tee ah vt crac etree! I. georgei Watling and Holman
— Pleonite 3 smooth, without keel, dorsal processes short, meet to

FORMA IVE eee ee ne en ee I. bransfieldi K. H. Barnard
6. Antenna 1, article 1, distal tooth extending past article 3; lateral

head sinus broadly rounded ........... G. sexdentata (Schellenberg)
— Antenna 1, article 1, distal tooth not extending past end of article

2; lateral head sinus narrow ............ G. macrops K. H. Barnard
7. Paired dorsal processes on pereonites 6 and 7 and pleonites 1-

Bit. Sit Re a dis ORE es erin 2 I. serrata (Schellenberg)
— Paired dorsal processes on pereonite 7 and pleonites 1-3 ........ 8
8. Coxa 7 posterior margin produced as an elongate process; pereopod

7 basis with 2 posterior teeth ...... I. acuticoxa Watling and Holman

— Coxa 7 smoothly rounded behind, at most with small tooth; per-
eopod 7 basis smoothly rounded with single small tooth at postero-

distalvcorner- 3.004 Ee a See A ee 9
9. Prominent distal tooth on antenna 1 peduncle article 1 extending

well past third peduncletanticle 224m... 240. 51 oe ae eee 10
— Teeth on antenna 1 peduncle article 1 not extending past third

article ee. FN A See PERRET TSI ee ese eed er 12

10. Coxa | rectangular, smoothly rounded anterodistally ...........
tease Nib YER me abate ES 8 G. mandibularis K. H. Barnard

— Coxa 1 not smoothly rounded, tapering distally, often bi-
GeMlate Vee oo. ete cys cue w Siete HAO ROPES Ce eee OE eee 11

VOLUME 94, NUMBER 1 187

11. Lateral head sinus a well rounded notch; mandible smooth;

(UDC prsnlanesttayle tose dns eo teal MPRA OP AMOR tat rag G. barnardi Thurston
— Lateral head sinus narrow; subacute mandible incisor multidentate,
Ching et eh nhc es ee ae ee ee I. margueritei Chevreux

12. Paired dorsal teeth form a distinct **V,’’ mid-dorsal keel present on

pleonites 1-3, additional upright keel on urosomite 1 ...........
er ee ries Pe ee eee Leclair: Barnard

— Paired dorsal teeth meet to form a ‘‘U,’’ no mid-dorsal keel
[SIRORSITE A gies Sic toc ee coe A ecient fee Sn Aaa etree. (2 Jc) Saunas ae ae 13

13. Coxa 1 distally tapering to an acute or subacute tip .............

mE COxa| LOUNnGed antenodistally "s. Feuve.. 026. ae ee ees he el ed 14
(4eeDornsal processes short, stubby; Upper lip entite =.7.-2,.----5---
oo a LOR AGE aS ON Se octet rnae a: I. microdentata (Schellenberg)
— Dorsal processes elongate, slender; upper lip incised ...........
RR ee er ee A ees MAS elon le I. cyclogena K. H. Barnard

Gnathiphimedia barnardi Thurston 1974
Fig. 3

Gnathiphimedia barnardi Thurston, 1974:15, figs. 3, 4.

Material—Eltanin Cruise 6, Sta. 410, 31 Dec. 1962, 61°18—20’S, 56°09-
10’W, 220-240 m, 1 a.

Diagnosis.—Body with paired dorsal processes on pereonite 7 and pleo-
nites 1-3; head, lateral sinus broadly rounded; antenna | peduncle article
2 dorsal tooth not extending beyond article 3; coxa | distally truncate; coxa
7 posterodistal corner slightly produced.

Description.—The following supplements the description of Thurston
(1974). Antennule with minute uniarticulate accessory flagellum; mandible
palp article 3 heavily setose, setae increasing in length distally; maxilliped
with minute article 4 concealed by hood-like extension of article 3; coxa 7
posterodistal corner slightly produced.

Distribution.—South Georgia, South Shetland Islands, 26-250 m.

Remarks.—Aside from the additions to Thurston’s description noted
above, the specimen from the South Shetland Islands differs from his ma-
terial as follows: the mandible palp and maxilla 1 inner plate and palp are
more setose; dorsal processes on pleon somite 3 are not as strong; upper
tooth on epimeral plate 3 is more dorsally positioned. Superficially, G. bar-
nardi resembles I[phimediella margueritei, especially with respect to pro-
cesses on the body and antennules. The two species can be distinguished
on the basis of mouthparts, head sinus (broad in G. barnardi) and coxa 6
(posteriorly acute in J. margueritei).

188 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Ss w

SIN \

SOC

Fig. 3. Gnathiphimedia barnardi, Eltanin Sta. 410: a, Antenna 1 peduncle articles 2, 3 and
accessory flagellum; b, Mandible; c, Maxilla 1; d, Distal end of maxilliped palp article 3 and
minute article 4; e, Coxae 6 and 7; f, Epimeral plates 2 and 3; g, Telson.

Gnathiphimedia macrops K. H. Barnard, 1932

Gnathiphimedia macrops K. H. Barnard, 1932:122, fig. 68.—Nicholls,
1938:78, fig. 41.—Thurston, 1974:13, fig. 2b.

Gnathiphimedia sexdentata.—K. H. Barnard, 1932:122 (in part).

Iphimediella bransfieldi K. H. Barnard, 1932:119 (in part).

Iphimediella discoveryi Watling and Holman, 1980:635.

VOLUME 94, NUMBER 1 189

Material.—Discovery Sta. 123, 15 Dec. 1926, off mouth of Cumberland
Bay, South Georgia, 230-250 m, 1 2 with eggs; Sta. 148, 9 Jan. 1927, off
Cape Saunders, South Georgia, 132-148 m, | 2 non-setose brood plates;
Sta. 175, 2 Mar. 1927, 63°17'S, 59°48’W, 200 m, | 2 with eggs.

Diagnosis. —Body with short, paired dorsal processes on pereonite 7,
pleonites 1 and 2; antenna | peduncle article | ventrodistal tooth extending
only to middle article 3; head lateral sinus bounded below by acute tooth,
bluntly rounded above; coxa | distally tapering and multidentate ventrally.

Description.—The following supplements the description given by K. H.
Barnard (1932) and Nicholls (1938). Body with 3 pairs of short processes
meeting dorsally in a narrow “‘U”’ shape. Upper lip entire. Mandible short
and stout but incisor smoothly dentate, not extremely thickened; molar
setose; palp article 3 only slightly shorter than article 2; accessory plate
present on both mandibles, better developed on left. Gnathopods | and 2
chelate: carpus and propodus short in gnathopod 1, carpus slightly longer
than propodus; gnathopod 2 carpus and propodus subequal. Pereopod
5 basis only slightly expanded, anterior and posterior margins parallel,
posterodistal corner produced into small acute point.

Distribution.—Adelie Coast and South Georgia, 120-540 m.

Remarks.—TYhe mandible of this species is not as heavily chitinized and
smooth as that found in other species of Gnathiphimedia. This, along with
some variation in length of the dorsal spines with body size, have probably
been the primary reasons for the confusion of this species with G. sexden-
tata and I. bransfieldi and the designation of one specimen as a new species,
I. discoveryi (Watling and Holman, 1980). Smaller specimens (from Sta. 123
and 148) carried proportionately longer dorsal teeth than did the larger fe-
male from Sta. 175. G. macrops can be distinguished from the only other
species of Gnathiphimedia with 3 pairs of dorsal processes, G. sexdentata,
by the short tooth on antenna | article | and by the shape of the lateral head
sinus. For a summary of the distinctions between J. bransfieldi and G.
macrops see the discussion concerning the differences between the former
and J. discoveryi in Watling and Holman (1980).

Gnathiphimedia sexdentata (Schellenberg 1926)

Diagnosis.—Body with paired dorsal processes on pereonite 7, pleonites
1 and 2; head lateral sinus broadly rounded; antenna | peduncle article |
ventrodistal tooth extending beyond peduncle article 3; coxa | distally ta-
pering, generally bidentate.

Gnathiphimedia sexdentata sexdentata, new subspecies
Figs. 4-6

Iphimediella sexdentata Schellenberg, 1926:331.
Iphimedia pacifica.—Walker, 1907 (part):27.—Chevreux, 1913:118.

190 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Gnathiphimedia sexdentata sexdentata, Terra Nova Sta. 8 male: a, Body, side view;
b, Lower lip. Female: c, Head side view; d, Head front view antennules and antennae removed;

e, Upper lip; f, Lower lip; g, Epimeral plates 2 and 3.

Gnathiphimedia pacifica.—K. H. Barnard, 1930:353, 449, fig. 27.
Gnathiphimedia sexdentata.—K. H. Barnard, 1932:122 (part).—Nicholls,

1938:77, fig. 40.—Thurston, 1974:13, fig. 2A.
Material.—Terra Nova Sta. 8 (2339) McMurdo Sound, 256 m, 2 66, 3

VOLUME 94, NUMBER 1 19]

ce

30o

)

os”

Fig. 5. Gnathiphimedia sexdentata sexdentata, Terra Nova Sta. 8, male: a, Antenna 1; b,
Mandible palp; c, Maxilla 2. Female: d, Antenna 1; e, Mandible; f, Maxilla 1; g, Maxilliped.

2°. Eltanin Cruise 6, Sta. 418, 2 Jan. 1963, 62°39-40’S, 56°08—-10’W, 311-
426 m, 2 juveniles; Cruise 51, Sta. 5769, 77°09.1'S, 158°59.6—159°00.2'’W,
344-357 m, 2 2 2, 1 juvenile.

Diagnosis.—Dorsal processes long, extending to posterior margin of fol-
lowing somite, projecting outward and slightly upward from body.

192 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

S WY

Fig. 6. Gnathiphimedia sexdentata sexdentata, Terra Nova Sta. 8, female: a, Pereopod 7
basis; b, Telson 21 mm specimen; c, Telson 22 mm specimen: d, Telson 28 mm specimen.

Description.—The following supplements the description given by K. H.
Barnard (1930). Antenna 1 peduncle article 2 ventrodistal tooth of variable
length, occasionally extending beyond peduncle article 3; accessory flagel-
lum uniarticulate. Upper lip lower margin entire to sinuous. Mandible in-
cisor thick, smooth; molar a short, broad-based cone, not as figured for G.
mandibularis (K. H. Barnard, 1930); accessory plates on right and left man-
dibles, that on right mandible minute, often broken; palp articles 2 and 3
subequal. Lower lip, distal apices vary from narrowly to broadly rounded.
Maxilla | palp article 2 armed along distal half of medial margin; outer plate
distally tapering; inner plate sub-acute, armed with approximately 10 plu-
mose setae. Maxilla 2 inner plate broadened distally. Maxilliped palp article
4 minute, not covered by hood-like extension of article 3, article 1 longer

VOLUME 94, NUMBER | 193

than article 3, article 2 not produced along article 3; outer plate pointed
distally, ovate; inner plate narrow, distally truncate.

Distribution.—Davis Sea, Adelie Coast, Ross Sea, Palmer Peninsula,
South Shetland Islands, 9—720 m.

Remarks.—Our examination of 5 British Museum specimens has dem-
onstrated this species to be extremely variable with regard to the following
features: antenna | peduncle article 2 distal tooth extends to or beyond
article 3; head sinus broad or narrow; mandible incisor of variable thickness
and degree of denticulation; coxa | narrowly or widely bidentate; maxilliped
palp article 2 equal to or shorter than article 1; pleurae on pereonites 3—7
lie flat against the body or project postero-laterally as acute processes. The
only constant character that separates G. s. sexdentata from G. s. incerta
appears to be the length of the dorsal body processes on pereonite 7 and
pleonites | and 2.

Gnathiphimedia sexdentata sexdentata, juvenile

Material.—Eltanin Cruise 6, Sta. 418, 2 Jan. 1963, 62°39-40’S, 56°08-
10’W, 311-426 m, 2 juveniles (6 mm length).

Remarks.—These specimens look very much like the adults described
above with the exception of the mandible which is much more slender.
Thus, these individuals are listed separately here in the event future workers
find the narrow mandible incisor not to be a juvenile character but rather
a feature indicative of a distinct species.

Gnathiphimedia sexdentata incerta, new subspecies
Figs. 7, 8

Gnathiphimedia incerta Bellan-Santini, 1972:170, figs. 2, 3.

Material.—Eltanin Cruise 6, Sta. 418, 2 Jan. 1963, 62°39-40'S, 56°08-—
10’W, 311-426 m, | 3; Cruise 12, Sta. 1003, 15 Mar. 1964, 62°41’S, 54°43’W,
210-220 m, 1 ¢, 1 2, 1 juvenile; Cruise 32, Sta. 1995, 10 Jan. 1968, 72°04’S,
172°38'E, 342-360 m, 2 juveniles.

Diagnosis.—Dorsal processes short, each pair projecting along contour
of body but not reaching to posterior margin of following somite.

Distribution.—Adelie Coast, South Shetland Islands, 60—426 m.

Remarks.—The morphological variation exhibited by these specimens
was similar to that listed for G. sexdentata sexdentata. Because of the
overlap in morphology for nearly all characters, G. incerta could not reliably
be distinguished from G. sexdentata other than by the length of the dorsal
body processes. Further study of many specimens will be required to fully
document the variability of the G. sexdentata complex.

194 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 7. Gnathiphimedia sexdentata incerta male. Eltanin Sta. 1003: a, Body side view; b,
Antenna 1. Male Eltanin Sta. 418: c, Right mandible. Female Eltanin Sta. 1003: d, Antenna
1; e, Coxae 1-3; f, Pleosome. Juvenile Eltanin Sta. 1003: g, Left mandible; h, Right mandible.

Iphimedia Rathke

Iphimedia Rathke, 1843:85.
Panoploea Thomson, 1880:2.—Karaman and J. L. Barnard, 1979:110.
Cypsiphimedia K. H. Barnard, 1955:87.—Watling and Holman, 1980:615.

Type-species.—l. obesa Rathke, 1843.
Diagnosis.—Upper lip entire or slightly emarginate; maxilla 1 palp biar-

VOLUME 94, NUMBER I 195

Fig. 8. Gnathiphimedia sexdentata incerta male. Eltanin Sta. 1003: a, Lower lip; b, Max-
illiped; c, Telson. Male Eltanin Sta. 418: d, Mandible palp; e, Telson. Female Eltanin Sta.
1003: f, Mandible palp.

ticulate, of variable length; maxilliped palp exceeding outer plate, 3-artic-
ulate, article 2 produced medially along article 3; gnathopod | chelate;
gnathopod 2 chelate or subchelate; telson variably emarginate or incised.

Remarks.—The above synonymy and current list of accepted species for
this genus is discussed in detail in Watling and Holman (1980).

Iphimedia joubini (Chevreux 1912)

Panoploea joubini Chevreux, 1912:212; 1913:114, figs. 19-21.—Nicholls,
1938:64, fig. 33 (var. bidentata).—Bellan-Santini, 1972:175, pl. 5.

Material.—Eltanin Cruise 6, Sta. 418, 2 Jan. 1963, 62°39-40’S, 56°08-
10’W, 311-426 m, | 2; Cruise 12, Sta. 1003, 1S Mar. 1964, 64°41’S, 54°43’'W,
210-220 m, 3 dd, 1 juvenile. Hero Cruise 731, Sta. 1946, 11 Mar. 1973,
64°52—53'S, 62°52—-53'W, 264-272 m, 1 juvenile. Burton Island Cruise 592,
Sta. 5, 29 Jan. 1958, 66°32.9’S, 92°54’E, 1 juvenile.

Diagnosis.—Body with elongate, paired dorsal processes on pereonite 7
and pleonites 1 and 2, with mid-dorsal keel on pleonite 3 extended poste-

196 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

riorly as acute process; pereopod 7 basis posterior margin excavate distally,
coxa 7 with single posterodistal tooth; coxa 5 posterior margin produced
as strong point.

Distribution.—Around most of the Antarctic Continent, from Davis Sea
to Adelie Coast, through western Antarctic to Palmer Archipelago and
South Shetland Islands, 45-540 m.

Remarks.—The specimens examined fit the original description of Chev-
reux very well, and thus would be assigned to the subspecies joubini as
diagnosed by Nicholls (1938).

Iphimedia multidentata (Schellenberg 1931)
Fig. 9

Panoploea multidentata Schellenberg, 1931:117, fig. 63.

Material.—Hero Cruise 715, Sta. 894, 2 Nov. 1971, 54°55’S, 64°18-20’W,
263-285 m, | o.

Diagnosis. —Body with paired dorsal processes on pereonite 7 and pleo-
nites | and 2; pleonite 3 with dorsal keel; pereopod 7 basis with 4 large teeth
on posterior margin; coxa 7 with 2 teeth on posterior margin.

Description.—The following supplements the description of Schellenberg
(1931). Head, anteroventral corner an acute process. Antenna | peduncle
article 1 with ventrodistal tooth extending to end of article 2; peduncle
article 2 with distal tooth dorsally, extending to end of article 3. Mandible
long and slender; incisor narrow, smooth; accessory plates on both man-
dibles, that on right mandible bidentate; palp article 3 half the length of
article 2. Maxilla 1 palp biarticulate, shorter than outer plate, article 2 twice
the length of article 1; inner plate not seen. Maxilliped palp 3-articulate,
article 2 strongly produced medially along article 3, the latter blunt distally;
outer and inner plates distally tapering. Gnathopod 1 chelate, dactyl tip
overhanging end of propodus fixed finger, each with a long plumose seta;
basis attached to coxa on inside of anterior margin and is directed forwards.

Distribution.—Falkland Islands and Magellanic area, 5—285 m.

Iphimediella Chevreux

Iphimediella Chevreux, 1911.
Pariphimediella Schellenberg, 1931 (part).—Watling and Holman, 1980.

Type-species.—l. margueritei Chevreux, 1912.

Diagnosis (modified from J. L. Barnard, 1969).—Antenna | accessory
flagellum uniarticulate; epistome not broad, less than 3 times as wide as
high; upper lip entire or weakly incised; mandible incisor elongate, not ex-
cessively thickened or broadened, oriented to cut in transverse plane; lower
lip without inner lobes, distal apices generally subacute; maxilla | palp bi-

VOLUME 94, NUMBER 1 197

Fig. 9. Iphimedia multidentata, Hero Sta. 894: a, Head and antenna | peduncle; b, Body
side view; c, Right mandible; d, Mandible palp; e, Maxilla 1. inner plate missing; f, Maxilliped;
g, Gnathopod 1; h, Gnathopod 1 propodus and dactylus; i, Coxa | inside showing position of
attachment to basis.

articulate, reaching end of outer plate; maxilliped palp 4-articulate, article
2 scarcely or not produced; gnathopods | and 2 chelate; telson slightly cleft.

Remarks.—The above synonymy, list of accepted species, and signifi-
cance of mandible incisor orientation are discussed in detail in Watling and
Holman (1980).

198 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

\
(/

Fig. 10. Iphimedielia margueritei, Eltanin Sta. 1003: a, Mandible; b, Mandible palp; c,
Maxilla 1; d, Maxilliped; e, Maxilliped palp articles 3 and 4; f, Coxa 1; g, Pereopod 6 coxa and
basis.

Iphimediella margueritei Chevreux 1912
Fig. 10

Iphimediella margueritei Chevreux, 1912:213.—Chevreux, 1913:120, figs.
22—24.—K. H. Barnard, 1930:348, fig. 22.
I. margueritei var. acuta Nicholls, 1938:69, fig. 36.

VOLUME 94, NUMBER I 199

Material.—Eltanin Cruise 12, Sta. 1003, 15 Mar. 1964, 62°41’S, 54°43’W,
PNO= 220 sm. 2aGner

Diagnosis.—Body with paired, broad-based, smooth dorsal processes on
pereonite 7 and pleonites 1-3; coxa | distally bidenticulate; antenna | pe-
duncle article 1 ventrodistal tooth extending beyond peduncle article 3; head
lateral margin with deep notch bounded above and below by acute pro-
cesses.

Description.—The following supplements the description given by Chev-
reux (1913). Antenna 1 with minute accessory flagellum. Mandible incisor
in smaller specimens multidenticulate. Maxilla 1 palp slightly longer than
outer plate, article 2 broadened, armed with plumose setae along distal half
of medial margin; outer plate distal margin tapering, armed with distal group
of strong spines, proximal margin with fine setae; inner plate subovate,
short, with few setae. Maxilliped palp article 4 small, partly covered by
hood-like extension of article 3; palp article 1 longer than article 3, article
2 slightly broadened; outer plate distal margin armed with few widely spaced
setae. Coxa | distally tapering, bidenticulate, teeth widely separated. Per-
eopods 6 and 7 coxae pointed posteriorly; bases posterior margin serrate.

Distribution.—Adelie Coast, Ross Sea, Antarctic Peninsula, Bransfield
Strait, Shag Rocks, South Georgia, South Sandwich Islands, 10-720 m.

Remarks.—The specimens described here differ only slightly from the
description given by Chevreux (1913). Some of these differences may be
related to the size of the individual; however it appears that the mandible
incisor becomes more robust and smooth as it wears. In both individuals
examined, the newly formed incisor, which could be seen through the old
exoskeleton, was finely dentate.

Iphimediella octodentata (Nicholls 1938)
Pariphimediella octodentata Nicholls, 1938:75, fig. 39.

Material.—Aurora, Sta. 2, 28 Dec. 1913, Commonwealth Bay, 527-550
m. (Holotype, Australian Museum #P18720.)

Diagnosis.—Body with paired dorsal processes on pereonites 6 and 7 and
pleonites 1 and 2, pleonite 3 with single, strong dorsal process; paired pro-
cesses noticeably serrate along upper margin.

Distribution.—Commonwealth Bay, 527-550 m.

Remarks.—Referring to I. octodentata, Nicholls (1938:75) stated ‘‘As
seen in profile this is scarcely to be distinguished from P. serrata Schell.
from which, however it differs in that the high median tooth on pleon 3 of
the latter is replaced in this present species by a pair of dorsal spines.’’ This
statement was interpreted to mean that while J. octodentata carried paired
dorsal processes on pleonite 3, Nicholls believed J. serrata carried only a
single tooth. As will be pointed out below J. serrata does, in fact, have

200 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

SS
ll

=
~
~
x
~
~

S

¢

Fig. 11. Iphimediella rigida, Eltanin Sta. 1002: a, Body side view; b, Anteroventral corner
of head and distal end coxa 1; c, Upper lip; d, Left mandible; e, Right mandible; f, Mandible
palp; g, Lower lip; h, Maxilla 1; 1, Maxilla 1 palp.

paired dorsal processes on pleonite 3, suggesting that J. octodentata and I.
serrata may be synonymous. Examination of the holotype of J. octodentata
showed that J. octodentata rather than I. serrata possessed the high median
tooth and the two species are therefore quite distinct.

Iphimediella rigida K. H. Barnard 1930
rics Pia

Iphimediella rigida K. H. Barnard, 1930:351, fig. 25.

VOLUME 94, NUMBER | 201

C

Fig. 12. Iphimediella rigida, Eltanin Sta. 1002: a, Maxilliped; b, Articles 3 and 4 of max-
illiped palp; c, Distal end gnathopod 1; d, Distal end gnathopod 2; e, Telson; f, Basis pereo-
pod 7.

Material.—Eltanin Cruise 12, Sta. 1002, 15 Mar. 1964, 64°40’S, 54°44—
45’W, 265 m, 1 2. Terra Nova Sta. 339, McMurdo Sound, 256 m, 1 6
(BMNH Cat. No. 1930. 8.1. 188).

Diagnosis. —Body with paired, short, smooth dorsal processes on per-
eonite 7 and pleonites 1-3; coxa | anteroventral corner broadly rounded;
antenna | peduncle article | medial ventrodistal tooth not extending beyond
article 3; head lateral margin with narrow notch bounded below by acute
process; maxilla 1 outer plate and maxilliped inner plate broadened.

Description.—The following supplements the short description given by
K. H. Barnard (1930). Head lateral margin with narrow notch bounded
above by a lobe bearing a short point, below by an acute process. Urosomite
1 with strong middorsal carina. Coxa | anteroventral corner rounded, ven-
tral margin slightly excavate. Coxa 2 and 3 anterior margins rounded, ta-
pering posterodistally to short points. Antenna | with minute accessory
flagellum. Upper lip slightly excavate. Mandible distally tapering, incisor
multidentate; molar conical without setae; accessory cutting plate multiden-
tate on right side, with fewer teeth on left side. Lower lip slightly emarginate
distally, inner lobes absent. Maxilla 1 palp article 2 with setae along three-
fourths of inner margin; outer plate broad, rhomboidal; inner plate short,
subtriangular. Maxilla 2 inner and outer plates broadened distally. Maxil-
liped palp article 4 minute, covered by hood-like extension of article 3; outer
plate narrow, armed distally with short, plumose and non-plumose setae;

202 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

inner plate truncate distally, as broad as outer plate. Gnathopod 1 dactyl
overhangs fixed finger of propodus, scarcely setose. Gnathopod 2 dactyl
shorter than fixed finger of propodus, the latter elongate and heavily setose.
Pereopod 7, basis posterior margin slightly serrate. Telson shallowly cleft.

Distribution.—McMurdo Sound, Bransfield Straits, 210—256 m.

Remarks.—This species is distinctive in the genus in its possession of a
strongly tapering mandible, broadened maxillae and maxilliped, and carinae
on pleonites 1—3 and urosomite |. The ovigerous female obtained from the
Eltanin material does not differ perceptibly from the male K. H. Barnard
described from McMurdo Sound.

Iphimediella serrata (Schellenberg 1926)
Fig. 13

Iphimedia serrata Schellenberg, 1926:328, fig. 43.
Pariphimediella serrata.—Schellenberg, 1931:121.

Material.—Eltanin Cruise 6, Sta. 418, 2 Jan. 1963, 62°39-40’S, 56°08-
10’W, 311-426 m, | juvenile.

Diagnosis. —Body with paired dorsally-serrate dorsal processes on per-
eonites 6 and 7 and pleonites 1—3; coxa | ventral margin serrate; head lateral
margin with broad, shallow notch.

Description.—The following description supplements that given by Schel-
lenberg (1926). Right mandible with multidenticulate accessory plate; molar
small, conical. Lower lip distally narrow. Maxilla 1 palp article 2 with setae
only on distal margin; outer plate distally tapering; inner plate short, acute,
with few setae on medial margin. Maxilla 2 outer plate distal setae nearly
as long as article. Maxilliped palp article 4 minute, covered by hood-like
extension of article 3; palp articles 1-3 subequal in length. Gnathopod 1
dactyl and fixed finger of propodus armed with long setae bearing recurved
setules. Gnathopod 2 fixed finger of propodus with row of minute setae.

Distribution.—Davis Sea (off Wilhelm II Coast), Bransfield Straits, 311-
426 m.

Remarks.—Much of the distinctiveness in the mouthparts of this species
is due to reduced or unusual setation. Since the mouthparts of an adult have
not been examined, it is difficult to determine whether these differences are
size-dependent. The form of the dorsal body processes distinguish J. serrata
from all other [phimediella species. Our finding of this species in the Brans-
field Straits extends the known distribution of this species to the opposite
side of Antarctica.

Maxilliphimedia K. H. Barnard
Maxilliphimedia K. H. Barnard, 1930:355.

VOLUME 94, NUMBER 1 203

hip
pu

Fig. 13. Iphimediella serrata, Eltanin Sta. 418: a, Mandible; b, Mandible palp; c, Lower
lip; d, Maxilla 1; e, Maxilla 2; f, Maxilliped; g, Gnathopod | tip; h, Gnathopod 2 tip.

Type-species.—Maxilliphimedia longipes (Walker 1906).

Diagnosis.—Epistome moderately broadened; upper lip broad, asymmet-
rically incised; mandible incisor broad but flattened, cutting edge multi-den-
tate, oriented to cut in frontal plane; lower lip without inner lobes; maxilla
1 palp biarticulate, article 2 greatly expanded; maxilla 2 inner and outer

204 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Sot eae

aoa ¢

SAMO»,
WON . ANAM ; ; =

Fig. 14. Maxilliphimedia longipes, Eltanin Sta. 1002: a, Gnathopod 1; b, Gnathopod 1| tip;
c, Gnathopod 2; d, Gnathopod 2 tip; e, Basis pereopod 7; f, Telson.

plates obovate; maxilliped palp 3-articulate, article 2 expanded but not pro-
duced along article 3; gnathopods chelate.

Maxilliphimedia longipes (Walker 1906)
Fig. 14

Iphimedia longipes Walker, 1906:151; 1907:29, pl. 9, fig. 17.
Maxilliphimedia longipes.—K. H. Barnard, 1930:355, fig. 28.

Material.—Eltanin Cruise 12, Sta. 997, 14 Mar. 1964, 61°44—-46’S, 55°54—
56'W, 769 m, 1 2 ovigerous; Cruise 12, Sta. 1002, 15 Mar. 1964, 62°40’S,
54°44-45'W, 265m, 14,1 2°.

Diagnosis.—As for genus.

Description.—The following supplements the description of K. H. Bar-
nard (1930). Gnathopod 1 chelate, basis with long setae along distal part of
lateral margin; article 6 longer than 5; fixed finger of propodus with several
hairs, partly covering dactyl. Gnathopod 2 chelate; article 6 longer than 5,
both articles with dense covering of setae; fixed finger of propodus longer
and thicker than dactyl, apically rounded. Pereopod 7 basis posterior margin
slightly concave distally.

Distribution.—Ross Sea to Bransfield Strait, 100—769 m.

Remarks.—This record extends the known range of the species from the
Ross Sea region to the tip of the Antarctic Peninsula.

Odius Lilljeborg

Otus Bate, 1862:125.
Odius Lilljeborg, 1865:19.—Stebbing, 1906:210.

VOLUME 94, NUMBER 1 205

Type-species.—Otus carinatus Bate, 1862 (original designation).

Diagnosis (emended from J. L. Barnard, 1969).—Upper lip narrow, in-
cised; mandible with narrow, dentate apex, molar prominent; palp of maxilla
1 uniarticulate, of varying length; maxilliped palp 4-articulate, article 2
slightly expanded but not produced along article 3; gnathopod 1 slender,
chelate; gnathopod 2 stout, subchelate; telson notched.

Odius antarcticus, new species
Figs. 15-17

Material.—Eltanin Cruise 12, Sta. 1003, 15 Mar. 1964, 62°41’S, 54°43’W,
210-220 m, 3 ovigerous 2 2, 5 mm maximum size.

Diagnosis.—Pleonites 1—3 and urosomite 1 dorsally smooth, without up-
right teeth; pleonite 3 posterior margin slightly gibbous; pereopods 5—7 bases
smoothly rounded behind without posterior teeth; eye small, not reniform;
maxilla 1 palp uniarticulate, long, reaching end of outer plate; maxilliped
palp 4-articulate, article 4 strongly reduced, only about '/10 length of article
3; telson visibly cleft.

Description.—Body smooth dorsally, pleonite 3 slightly gibbous. Rostrum
well-developed, extending past antenna | peduncle article 1 distal margin.
Coxa 1-3 longer than wide. Coxa 4 ventral margin straight. Eye small, not
reniform. Antenna | shorter than antenna 2; lacking accessory flagellum.
Upper lip longer than wide; distal margin asymmetrically incised. Mandible
incisor multidentate; accessory plate present on left side; molar prominent,
setose; strong spine row present between incisor and molar; on left man-
dible, spines closest to incisor widened distally and become multidentate,
similar in shape to the accessory plate; palp 3-articulate, articles 2 and 3
subequal in length. Lower lip notched along midline; apices narrow, setose.
Maxilla | with uniarticulate palp exceeding length of outer plate; outer plate
carrying short, heavy spines distally, densely setose proximally; inner plate
with 3 plumose setae distally. Maxilla 2 outer plate narrower than inner;
setae on outer plate twice length of those on inner. Maxilliped palp 4-artic-
ulate, article 2 slightly expanded medially, article 4 small. Gnathopod 1
chelate, articles 5 and 6 subequal. Gnathopod 2 strongly subchelate, palm
border serrate. Pereopods 5-7 similar, posterior margin bases smooth, per-
eopod 7 basis slightly more elongate. All uropods biramous; uropod | inner
and outer rami subequal; uropod 2 inner ramus longer and wider than outer;
uropod 3 inner ramus approximately 1.5 x length of outer; telson longer
than wide, cleft %.

Distribution.—Known only from type-locality.

Remarks.—Three other species are included within the genus Odius: O.
carinatus Bate 1852; O. kelleri Bruggen, 1907; O. cassigerus Gurjanova,
1972. All 3 of the above species have a minute uniarticulate palp on maxilla

206 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 15. Odius antarcticus, Eltanin Sta. 1003: a, Body side view; b, Antenna 1; c, Antenna
2; d, Upper lip; e, Mandible; f, Mandible incisor and spine row; g, Mandible palp; h, Lower
lip; 1, Maxilla 1.

VOLUME 94, NUMBER | 207

Fig. 16. Odius antarcticus, Eltanin Sta. 1003: a, Maxilla 2; b, Maxilliped; c, Gnathopod 1;
d, Gnathopod 1 tip; e, Gnathopod 2; f, Pereopod 5; g, Pereopod 6.

208 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 17. Odius antarcticus, Eltanin Sta. 1003: a, Pereopod 7 articles 2-4; b, Uropod 2; c,
Uropod 3; d, Telson.

1 whereas the palp is quite large for the new species described here. Simi-
larities in basic body shape and in most other mouthparts, especially the
mandibles, serve as unifying features, however, and the difference in palp
length was not considered to be outside the scope of the genus. O. antarc-
ticus iS also distinctive from all other species of Odius heretofore described
by its smooth body profile, small fourth article on the maxilliped palp,
smooth bases of the pereopods, and the degree to which the telson is cleft.

Etymology.—tThis is the first known species of Odius to be described
from the Antarctic.

Holotype. —USNM 181323; 2 2° paratypes USNM 181324.

Parapanoploea Nicholls
Parapanoploea Nicholls, 1938:65.

Type-species.—Parapanoploea oxygnathia Nicholls, 1938, p. 66.

Diagnosis (modified from J. L. Barnard, 1969).—Upper lip broad, faintly
emarginate; mandible incisor narrow, elongate; lower lip lobes not incised;
maxilla 1 palp biarticulate, reaching end of outer plate; maxilliped palp

VOLUME 94, NUMBER | 209

d

Fig. 18. Parapanoploea oxygnathia, Eltanin Sta. 5769: a, Head; b, Body side view; c,
Antenna | peduncle; d, Mandible; e, Maxilla 2; f, Gnathopod 1.

4-articulate, article 2 slightly expanded but not produced along article 3;
gnathopods | and 2 chelate; telson notched.

Parapanoploea oxygnathia Nicholls 1938
Fig. 18

Parapanoploea oxygnathia Nicholls, 1938:66, figs. 34, 35.

Material.—Eltanin Cruise 51, Sta. 5769, 9 Feb. 1972, 77°09.1'S, 158°59.6’—
159°00.2'W, 344-357 m, | juvenile.
Diagnosis.—Rostrum one-half of antenna 1 peduncle article 1;

210 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

distal tooth on antenna | peduncle article | extends at least to distal margin
of peduncle article 2; gnathopod 1 article 6 4/s the length of article 5; dorsal
processes on pereonite 7 and pleonites | and 2 directed posteriorly.

Description.—The following supplements the description of Nicholls
(1938). Body with paired dorsal processes, separated by wide U-shaped gap,
on pereonite 7 and pleonites 1 and 2; processes on pleonite | parallel with
dorsal margin of pleonite. Antenna 1 peduncle article 1 distal tooth extends
to distal margin peduncle article 2.

Distribution.—Davis Sea and Ross Sea, 216-357 m.

Remarks.—Since the specimen at hand is a juvenile, it is with some hes-
itation that we have assigned it to P. oxygnathia. While it agrees with
Nicholls’ (1938) description in terms of the distal tooth on antenna 1 pe-
duncle article 1, and the length of gnathopod | article 6, it otherwise was
very similar to P. longirostris Bellan-Santini (1972), who differentiated the
2 species using the following characters: (1) lateral margin of the head at
the level of the eye is sinuous in P. longirostris, simply rounded in P.
oxygnathia; (2) mandible incisor possesses 8 teeth in P. longirostris, 6 in
P. oxygnathia; (3) rostrum extends 4/s the length of antenna | peduncle
article 1 in P. longirostris, versus one-half in P. oxygnathia; (4) gnathopod
1 merus/carpus ratio is opposite in the 2 species; (5) uropod 3 rami equal in
P. longirostris, unequal in P. oxygnathia. Characters (1), (2), (3), and (4)
are slight differences which eventually may be found to be gradational (to
date, only 2 specimens of P. oxygnathia and | of P. longirostris are known).
Nicholls’ illustration ef uropod 3 suggests that the inequality of the rami is
due to the outer ramus being broken. Our specimen was missing both rami
of uropod 3 so this could not be checked. Thus, it appears for the present
that the real differences between these 2 species are: (1) the presence of the
long distal tooth on antenna 1 peduncle article 1 in P. oxygnathia; and (2)
the backwardly-directed paired dorsal processes on pleonites | and 2 in P.
oxygnathia versus the upwardly-directed paired dorsal processes on pleo-
nites 1 and 2 in P. longirostris. Bellan-Santini does not figure the paired
dorsal processes on pereonite 7 in P. longirostris, therefore no comparison
of this feature can be made with P. oxygnathia.

Paramphithoidae
Epimeria Costa

Epimeria Costa in Hope, 1851:46.
Pseudepimeria Chevreux, 1911:1167.
Subepimeria Bellan-Santini, 1972:225.—Watling and Holman, 1980.

Type-species.—Gammarus corniger J. C. Fabricius, 1779.
Diagnosis (emended from J. L. Barnard, 1969).—Rudimentary accessory
flagellum; mandible molar large, ridged; lower lip lacking inner lobes; max-

VOLUME 94, NUMBER 1 2a

le g

Fig. 19. Epimeria georgiana, Eltanin Sta. 1003: a, Body side view; b, Antenna 1; c, Lower
lip; d, Maxilliped: e, Maxilliped articles 3 and 4 with setae omitted; f, Gnathopod | articles
6, 7; g, Telson.

illiped palp 4-articulate; gnathopods simple to subchelate, dactyls often spi-
nose, much shorter than propodus; coxae 4—S together forming a more or
less crescentic curve below.

Epimeria georgiana Schellenberg 1931
Fig. 19

Epimeria georgiana Schellenberg, 1931:160.
Epimeria excisipes K. H. Barnard, 1932:174, figs. 104e, 106, 107 (new syn-
onymy).

Material.—Eltanin Cruise 6, Sta. 410, 31 Dec. 1962, 61°18—20’S, 56°09-
10’W, 220-240 m, | 2; Cruise 12, Sta. 1002, 15 Mar. 1964, 62°40’S, 54°44—
45'W, 265 m, 9 22 (6 with eggs, 1 with young), 1 6; Cruise 12, Sta.
1003, 15 Mar. 1964, 62°41’'S, 54°43’W, 210-220 m, 1 2 with eggs; Discovery
Sta. 42, 1 Apr. 1926, 120-204 m (Br. Mus. #1936. 11.2. 1551-1580); Cum-
berland Bay, South Georgia (54°11'S, 36°18’W), 5 June 1902, 252-310 m,
(Naturhistoriska Riksmuseet Type No. 673).

Diagnosis.—Body with dorsal carinae on at least pereonites 4—7 and ple-
onites 1-3; coxa 4 anteroventral margin slightly concave, not acutely point-
ed ventrally; coxa 5 not produced posteriorly; pereopods 5—7 bases, hind
margins with distal notch.

Description.—The following supplements the description of Schellenberg
(1931) and K. H. Barnard (1932). Antenna 1 with minute accessory flagel-

2)\ 72 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

lum; peduncle article 2 twice as long as wide. Lower lip apex subacute;
medial margin with dense cover of setae. Maxilliped palp article 2 longer
than articles 1 and 3, article 4 with many closely-spaced short spines on
inner margin. Gnathopod 1 subchelate; article 6 three-fourths as wide as
long, subequal to article 5; palm circular, finely serrate; dactyl inner margin
with many closely-spaced spines. Third epimeral plate posterodistal margin
with acute tooth, without accessory tooth. Telson cleft nearly one-third.

Distribution.—Palmer Archipelago, Bransfield Strait, South Shetland Is-
lands, South Georgia, 75-391 m.

Remarks.—K. H. Barnard (1932) suggested that E. excisipes was prob-
ably a synonym of E. georgiana Schellenberg (1931). We examined the
types of both species and agree with K. H. Barnard’s suggestion. J. L.
Barnard (1961) separated the 2 in his key to the species of Epimeria by the
presence of an accessory tooth above the lower posterior corner of epimeron
3 in E. excisipes. McCain (1971) suggested that coxa 4 was “‘broadly quad-
rate’ in E. excisipes and ‘“‘narrowly truncate”’ in E. georgiana. Neither of
the type specimens had an accessory tooth on epimeron 3, and their fourth
coxae could both be described as broadly quadrate.

Epimeria macrodonta Walker 1906
Fig. 20

Epimeria macrodonta Walker, 1906:16.—Walker, 1907:24, pl. 8, fig. 14.

Epimeria similis Chevreux, 1912:215; 1913:149, figs. 41-43.

Epimeria macrodonta forma macrodonta K. H. Barnard, 1930:372;
1932:172, fig. 105.

Epimeria macrodonta forma similis K. H. Barnard, 1930:372; 1932:172.

Material.—Eltanin Cruise 6, Sta. 410, 31 Dec. 1962, 61°18—20’S, 56°09-
10’W, 220-240 m, 1 ¢ (f. similis); Cruise 12, Sta. 1002, 15 Mar. 1964,
62°40’'S, 54°44—45’W, 265 m, 3 2 @ (f. similis); Cruise 12, Sta. 1003, 15 Mar.
1964, 62°41’S, 54°43’W, 210-220 m, 4 22 (3 ovigerous), 3 juveniles, 2
3 6 (f. similis). Hero Cruise 731, Sta. 1946, 11 Mar. 1973, 64°53’S, 62°53’W,
264-272 m, 1 @ (f. macrodonta).

Diagnosis.—Body with dorsal carinae on pleonites 1—3 and most pereon-
ites, but never on pereonite 2; urosomite 1 with mid-dorsal upright tooth;
coxa 5 acutely produced posteriorly; pereopods 5 and 6 bases posterior
margin without notch, posterodistal corners acutely produced, anterior and
posterior margins parallel.

Description.—The following supplements the description of Walker (1907)
and Chevreux (1913). Antenna | with minute accessory flagellum. Upper lip
asymmetrically incised. Mandible palp article 3 slightly longer than article
2, with proximal seta group on dorsal margin. Lower lip apex blunt, with
tuft of stiff setae on medial margin at apex. Maxilla 1 palp with stout setae

VOLUME 94, NUMBER | ZS

Uy
a a
}

\

NN

a

Fig. 20. Epimeria macrodonta, Eltanin Sta. 1002: a, Upper lip; b, Mandible palp; c, Lower
lip: d, Maxilla 1; e, Maxilliped; f, Maxilliped articles 3 and 4 with setae omitted; g, Gnatho-
pod 1.

at tip, grading into fine setae proximally along medial margin; inner plate
broader than outer with setae long and thick distally, shorter, thinner and
more plumose proximally. Maxilliped palp articles 1-3 subequal in length;
article 4 covered proximally by slight extension of third article, with several
short spines on medial margin. Gnathopod 1 subchelate; palm narrowly
rounded, finely serrate; dactyl with many short, closely-spaced spines on
inner margin; propodus linear.

Distribution.—Circum-Antarctic, 30—900 m.

Remarks.—The specimen from Station 410 is similar to the other speci-
mens of E. macrodonta f. similis except that it does not possess a carina
on pereonite | and its antenna | peduncle article 2 possesses a mid-ventral
tooth on the distal margin. This form is presently regarded as being aberrant
and not deserving of separate subspecies status.

Epimeria puncticulata Barnard 1930
Fig. 21

Epimeria puncticulata K. H. Barnard, 1930:376, fig. 42; 1932:175, fig. 104d.
Subepimeria geodesiae Bellan-Santini, 1972:225, figs. 33, 34 (new synonymy).

214 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

)

Dy

—

SS
SS
SS SS Sana

Fig. 21. Epimeria puncticulata, Eltanin Sta. 1003: a, Pleonites 1-3 dorsal view; b, Antenna
1; c, Mandible palp; d, Lower lip; e, Maxilla 1; f, Maxilliped; g, Maxilliped articles 3 and 4
with setae omitted; h, Gnathopod 1; i, Pereopod 5; j, Pereopod 6; k, Pereopod 7.

Material.—Eltanin Cruise 12, Sta. 1003, 15 Mar. 1964, 62°41’S, 54°43’W,
210-220 m, 1 2 with eggs; Cruise 32, Sta. 1995, 10 Jan. 1968, 72°04’S,
172°38'E, 360-342 m, | juvenile. Discovery 1936.11.2, 1581-1590 m, (British
Museum), South Georgia, 2 2°.

Diagnosis.—Body without dorsal carinae on pereonites, carinae present

VOLUME 94, NUMBER 1 Dal)

on pleonites 1-3; coxa 5 not produced posteriorly; pereopod 5 basis pos-
terodistal corner subacutely produced, hind margin without notch.

Description.—The following supplements the description of Barnard
(1930). Carinae on pleonites 1—3 forming a distinct keel, the second strongly
pointed posteriorly. Antenna | with minute, uniarticulate accessory flagel-
lum; peduncle article 2 as wide as long. Lower lip apically subacute, with
strong group of stiff setae on apex medial margin. Mandible palp article 3
as long as article 2, armed along entire ventral margin with setae, last 3
setae double the length of others. Maxilla 1 palp biarticulate, armed with
distal group of stout setae; inner plate apical setae stout, plumose. Maxil-
liped palp 4-articulate, articles 1-3 subequal in length; article 4 with few
spines along medial margin. Gnathopod | subchelate; articles 5 and 6 elon-
gate, slightly longer than article 6; dactyl with 6 spines on inner margin;
palm excavate near base of dactyl, proximal margin finely serrate. Pereopod
5 basis posterior margin excavate, posterodistal corner subacute. Pereopod
6 basis posterodistal corner rounded. Pereopod 7 basis broader than in pereo-
pods 5 and 6, hind margin convex. Telson cleft one-fourth.

Distribution.—Ross Sea, Antarctic Peninsula, South Georgia, 60-270 m.

Remarks.—Our specimen from Eltanin Sta. 1003 differed from the Dis-
covery material examined in lacking a small mid-dorsal tooth on the pos-
terior margin of pereonite 7; however, in all other respects they were indis-
tinguishable. Of the species in this genus which lack dorsal carinae on the
pereonites, the following 4 species form a group characterized by distally
broad 4th coxae and posterodistally blunt 5th coxae: E. geodesiae (Bellan-
Santini), E. monodon Stephenson, E. puncticulata K. H. Barnard and E.
robusta K. H. Barnard. The posteriorly notched bases on pereopods 5 and
6 readily separate E. robusta from this group. Pereopods 6 and 7 bases are
both posteriorly expanded in E. monodon whereas, in E. geodesiae and E.
puncticulata, only the basis of pereopod 7 is posteriorly expanded. We have
elected to synonymize E. geodesiae with E. puncticulata since the 2 differ
only in the presence or relative strength of a mid-dorsal carina on pleonite
1. We have seen some variability in this character as noted above, and as
well, K. H. Barnard (1916) found that for large specimens of E. semiarmata,
such carinae could become obsolete.

Metepimeria Schellenberg
Metepimeria Schellenberg, 1931:162.

Type-species.—Metepimeria acanthurus Schellenberg, 1931.

Diagnosis (emended from J. L. Barnard, 1969).—Antenna 1 with acces-
sory flagellum; mandibular molar large, ridged; lower lip without inner
lobes; maxilliped palp 3-articulate; gnathopods simple.

216 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 22. Metepimeria acanthurus, Hero Sta. 894: a, Antenna 1; b, Upper lip; c, Mandible;
d, Mandible palp; e, Lower lip; f, Maxilla 1; g, Maxilla 2; h, Maxilliped; 1, Gnathopod | (setae
omitted); j, Gnathopod 2 (setae omitted); k, Telson.

Metepimeria acanthurus Schellenberg 1931
Fig. 22

Metepimeria acanthurus Schellenberg, 1931:162, fig. 85, pl. 1, fig. G.
Epimeria acanthurus.—K. H. Barnard, 1932:176, figs. 104B, 108; pl. 1,
fig. 2.

Material.—Eltanin Cruise 9, Sta. 740, 18 Sept. 1963, 56°06—-07'S, 66°19-—
30’W, 384-494 m, | 2°. Hero Cruise 715, Sta. 894, 2 Nov. 1971, 54°54.8-
55'S, 64°18-20'W, 263-285 m, 1 2,936. William Scoresby Sta. 86, 3 Mar.
1927, 53°53'S, 60°34’W, 151-147 m, | & ovigerous.

Diagnosis.—As given for genus.

Description.—The following supplements the description of Schellenberg

VOLUME 94, NUMBER | 217

(1931) and K. H. Barnard (1932). Antenna 1 with minute accessory flagel-
lum, bearing single terminal seta. Upper lip shallowly notched. Mandible
incisor multidenticulate, left lacinia mobilis with 6 teeth; molar large, trit-
urative, with lateral and anterior setose ridges; palp articles 2 and 3 subequal
in length; palp article 3 with heavy cover of long setae on ventral margin,
article 2 with several setae on distal half of ventral margin. Lower lip without
inner lobes; medial margin near distal apex with group of short, blunt, stiff
setae. Maxilla 1 palp article 2 with setae along distal half of medial margin;
inner plate with long, thick, sparsely plumose setae. Maxilla 2 inner and
outer plates broadened distally. Maxilliped palp 3-articulate, palp article 2
slightly longer than article 3; palp article 3 with dense apical tuft of setae
arranged in 3 whorls; outer plate extends halfway along palp article 2.
Gnathopod | simple, articles 5 and 6 subequal in length, article 5 wider than
article 6; dactyl armed with series of short, stiff spines along inner margin.
Gnathopod 2 simple, articles 5 and 6 longer than in gnathopod 1, subequal
in length and width; dactyl armed with series of stout, stiff spines. Telson
shallowly cleft.

Distribution.—Magellanic area, Falkland Islands, 151-194 m.

Remarks.—This species is very similar to the species in the genus Epi-
meria, especially in the form of the mandible, lower lip and maxillae. The
absence of a fourth article on the maxilliped palp combined with the simple
gnathopods does not allow its inclusion in the genus Epimeria despite its
superficial resemblance to E. puncticulata (as noted by K. H. Barnard,
1952)

Uschakoviella Gurjanova
Uschakoviella Gurjanova, 1955:199.

Type-species—U. echinophora Gurjanova, 1955.

Diagnosis (as modified from J. L. Barnard, 1969).—Body covered with
articulated spines; accessory flagellum minute; lower lip lacking inner lobes;
mandible molar large, ridged; maxilliped palp 4-articulate; gnathopods
scarcely subchelate but palms transverse; telson cleft.

Uschakoviella echinophora Gurjanova 1955
Fig. 23

Uschakoviella echinophora Gurjanova, 1955:200, figs. 14, 15.
U. e. echinophora Gurjanova, 1955:203.

U. e. abyssalis Gurjanova, 1955:203, figs. 16-18.

U. echinophora.—Shoemaker, 1964:417, fig. 12.

Material.—Islas Orcadas Cruise 19, Sta. 14, 18 Mar. 1979, 59°48’S,
45°06'W, 1 juvenile.

218 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 23. Uschakoviella echinophora, Islas Orcadas Cruise 19, Sta. 14: a, Coxae 1-4; b,
Antenna 1; c, Lower lip; d, Maxilla 1; e, Telson.

Diagnosis.—See generic diagnosis.

Description.—The following supplements the description given by Gurja-
nova (1955) and Shoemaker (1964). Spines on body become slightly heavier
at posterior of body. Pleonite 3 has, in addition to single upright dorsal
tooth, a slightly more slender tooth at posterior margin. Antenna | accessory
flagellum minute. Upper lip incised. Maxilla 1, inner plate with 5 ‘‘velvety”’
setae distally; palp with only short spines. Lower lip with single prominent
seta at distomedial margin. Telson with 4 prominent dorsal spines, lobes
with minute grooves distally.

Distribution.—Boreal North Pacific, Antarctic, S54—2550 m.

Remarks.—Our specimen differs from those of Shoemaker (1964) and
Gurjanova (1955) in the possession of short spines on the palp of maxilla 1
instead of longer, tufted setae and in the presence of prominent spines on
the telson. The presence of the accessory flagellum and the additional small-
er tooth on pleonite 3 was probably overlooked in the earlier studies as both

VOLUME 94, NUMBER | INE)

these characters are difficult to see. The importance of the additional char-
acters 1S uncertain because of disagreements in the earlier descriptions.
Shoemaker described the upper lip of his specimens as broad, symmetrical
and not incised while Gurjanova figured an incised upper lip. Gurjanova
illustrated 2 distomedial spines on the lower lip whereas Shoemaker figured
only fine setae. Because this species has been reported so few times, as-
sessment of possible specific differences 1s difficult.

Stegocephalidae
Andaniotes Stebbing

Andaniotes Stebbing, 1897:30.
Metandania Stephensen, 1925:136.

Type-species.—Anonyx corpulentus Thomson, 1882.

Diagnosis (from J. L. Barnard, 1969).—Antenna | flagellar article | equal
to or shorter than peduncle; antenna 2 peduncle article 5 equal to or shorter
than article 4; mandible incisor not toothed; maxilla 1 palp uniarticulate;
maxilla 2 outer plate not geniculate or gaping; maxilliped palp article 2 not
produced; pereopod 5 article 2 slender; pereopods 6 and 7 article 2 broad;
pleonite 6 longer than uropod 3 peduncle.

Andaniotes corpulentus (Thomson 1882)
Fig. 24

Anonyx corpulentus Thomson, 1882:231, pl. 17, fig. 1.

Andania abyssorum Stebbing, 1888:739, pl. 37.

Andaniotes corpulentus.—Stebbing, 1897:31, pl. 8; 1906:96, fig. 21.—Schel-
lenberg, 1931:51.—J. L. Barnard, 1972:307 (key).

Material.—Eltanin Cruise 6, Sta. 410, 31 Dec. 1962, 61°18—20'S, 56°09-
10’W, 220-240 m, 2 individuals; Cruise 6, Sta. 428, 5 Jan. 1963, 62°39-41’S,
57°46—-51'W, 622-1120 m, 1 2 ovigerous; Cruise 9, Sta. 740, 18 Sept. 1963,
56°06—07'S, 66°19-30'W, 384-494 m, 1 &, 3 juveniles, 1 sex unknown;
Cruise 12, Sta. 1003, 15 Mar. 1964, 62°41’S, 54°43’W, 210-220 m, 1 individ-
ual; Cruise 12, Sta. 1079, 13 Apr. 1964, 61°25—26’S, 40°55’W, 593-598 m,
l 2.

Diagnosis.—Maxilla | palp not reaching end of outer plate; maxilliped
palp 4-articulate; pereopod 6 article 2 slightly expanded, posterior margin
distally rounded, article 2 more than twice width article 3.

Description.—The following supplements the description of Stebbing
(1888). Antenna 1 accessory flagellum uniarticulate. Epistome tall, usually
with small conical projection from ventral margin, one specimen (?male)
from Eltanin Sta. 410 with large, ventral, anteriorly-directed recurved

220 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 24. Andaniotes corpulentus, Eltanin Sta. 410: a, Head; b, Antenna 1; c, Lower lip; d,
Maxilla 1; e, Maxilla 2; f, Maxilliped; g, Pereopod 6; h, Pereopod 7; 1, Head with unusual
epistome.

tooth. Maxilla 1 more heavily setose than described by Stebbing. Maxilliped
inner plate truncate distally, inner corner produced as a tooth; palp article
1 more than twice as long as wide. Pereopod 6 basis hind margin slightly
convex. Pereopod 7 basis broadly rounded posteriorly.
Distribution.—Falkland Islands, Magellanic Area, South Shetland Is-

VOLUME 94, NUMBER | 221

HY, | ) |
Ny

Fig. 25.. Andaniotes linearis, Islas Orcadas Cruise 575, Sta. 91: a, Antenna 1; b, Antenna
2; c, Epistome and upper lip; d, Mandible; e, Lower lip; f, Maxilla 1; g, Maxilla 2.

lands, South Orkney Islands, Atlantic sector of Southern Ocean, and New
Zealand, low tide to 1120 m.

Andaniotes linearis K. H. Barnard 1932
Figs. 25, 26

Andaniotes linearis K. H. Barnard, 1932:80, fig. 36.—Nicholls, 1938:41,
ny, Alle

222 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ane

ni

y

Fig. 26. Andaniotes linearis, Islas Orcadas Cruise 575, Sta. 91: a, Maxilliped; b, Gnathopod
1; c, Gnathopod 2; d, Pereopod 5; e, Pereopod 6; f, Pereopod 7; g, Telson.

Material.—Eltanin Cruise 6, Sta. 410, 31 Dec. 1962, 61°18—20'S, 56°09—
10’W, 220-240 m, 7 2 2, 8 juveniles; Cruise 6, Sta. 428, 5 Jan. 1963, 62°39-
Al’S, 57°46-S1'W, 622-1120 m, 1 2 with young, | juvenile; Cruise 7, Sta.
558, 14 Mar. 1963, 51°58’—52°01'S, 56°38’ W, 646-845 m, 1 2°; Cruise 9, Sta.
684, 25 Aug. 1963, 54°55’S, 38°0S—07'’W, 595-677 m, 11 22, 1 3; Cruise 9,
Sta. 740; 18 Sept. 1963, 56°06—-07'S, 66°19-30'W, 384-494 m, 2 92, 1 sex

VOLUME 94, NUMBER 1 22S

unknown; Cruise 11, Sta. 977, 13 Feb. 1964, 52°32’S, 63°53'’W, 229 m, 1
2 ovigerous, 3 juveniles. Islas Orcadas Cruise 575, Sta. 82, 6 June 1975,
55°29.0'S, 35°20.5’W, 413-462 m, 1 juvenile; Cruise 575, Sta. 90, 7 June
1975, 54°50.6’S, 37°23.8’W, 223-227 m, 4 2° 2 (3 with eggs), 3 66, 1 juve-
nile: Cruise 575, Sta. 91, 7 June 1975, 55°00.6'S, 37°42.6’W, 494—501 m, 1
2 with eggs, 6 ¢6 64, 4 juveniles. Hero Cruise 715, Sta. 894, 2 Nov. 1971,
54°55'-54.8'S, 64°20-18’W, 263-285 m, 1 juvenile.

Diagnosis.—Maxilla | palp reaching outer plate; maxilliped palp 4-artic-
ulate; pereopod 6 article 2 rectilinear, twice as wide as article 3.

Description.—Antenna 1 accessory flagellum long but uniarticulate,
armed distally with a single long seta; main flagellum of 4 articles, first
article more than half length of peduncle. Antenna 2 peduncle article 4
longer than article 5. Epistome triangular, as long as basal width, with low
medial keel. Upper lip asymmetrically incised. Mandible greater than twice
as long as broad; molar conical. Lower lip distal apices elongate, densely
covered with setae, proximal apices short, subacute. Maxilla 1 palp reaching
end of outer plate; inner plate not as broad as outer plate. Maxilla 2 inner
plate 3 times width of outer plate at apex. Maxilliped palp 4-articulate; palp
article 1 less than twice as long as wide; outer and inner plates broad.
Gnathopod | propodus with dense covering of setae on distal half of dorsal
margin. Gnathopod 2 propodus linear, without well-defined palm. Pereopod
5 basis linear, as wide as article 3. Pereopod 6 basis rectilinear, twice width
article 3. Pereopod 7 basis hind margin broadly rounded, tapering distally.
Uropod 3 outer ramus biarticulate. Telson cleft one-half.

Distribution.—Adelie Coast, around western Antarctica to Palmer Ar-
chipelago, South Georgia and Falkland Islands, 81-1120 m.

Euandania Stebbing
Euandania Stebbing, 1899:206.

Type-species.—Andania gigantea Stebbing, 1888.

Diagnosis (from J. L. Barnard, 1969).—Mandibular incisor not toothed;
maxilla | palp uniarticulate; maxilla 2 outer plate not geniculate or gaping;
maxilliped palp article 2 not produced; pereopod 5 article 2 slender; per-
eopod 6 article 2 broad or slender; pereopod 7 article 2 broad; telson cleft
one-third; antenna | flagellar article 1 much longer than peduncle; pleonite
6 shorter than peduncle of uropod 3.

Euandania gigantea (Stebbing 1883)
Fig. 27a—d

Andania gigantea Stebbing, 1883:206; 1888:730, pl. 35.
Euandania gigantea.—Stebbing, 1899:206.—K. H. Barnard, 1932:80.

224 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 27. Euandania gigantea, Islas Orcadas Cruise 575, Sta. 6: a, Lower lip; b, Pereopod
7; c, Adult telson; d, Juvenile telson. Parandania boecki, Eltanin Sta. 466: e, Antenna 2.

Material.—Eltanin Cruise 9, Sta. 720, 7 Sept. 1963, 56°0S—07’'S, 34°00-
03'’W, 2818-2873 m, 1 juvenile; Cruise 10, Sta. 862, 21 Nov. 1963, 61°17-—
26'S, 78°55-57'W, 4575 m, 1 juvenile. /slas Orcadas Cruise 575, Sta. 6, 9
May 1975, 51°02.2'S, 42°47.6'W, 1480-1545 m, 1 @.

Diagnosis.—Antenna 1 flagellar article | much longer than peduncle; an-
tenna 2 peduncle article 5 longer than article 4; pleonite 6 shorter than
peduncle of uropod 3; pereopod 6 article 2 broad.

Description.—The following supplements the description of Stebbing
(1888). Lower lip medial margin with small protuberance near distal apex.
Pereopod 7 basis broad, tapering distally. Telson widely cleft.

Distribution.—Ross Sea, Prince Edward Islands, Indian and Pacific sec-
tors of Southern Ocean, Kermadec Trench, South and Northwest Atlantic
Oceans, bathypelagic, 0—4575 m.

Remarks.—Our specimens differed from that illustrated by Stebbing
(1888:35) in having pereopod 7 basis slightly more expanded. There ap-
peared to be some slight variation in the degree to which the telson was
cleft; that of one junveile was only slightly cleft.

Parandania Stebbing
Parandania Stebbing, 1899:206.

Type-species.—Andania boecki Stebbing, 1888.
Diagnosis (from J. L. Barnard, 1969).—Mandible incisor not toothed;

VOLUME 94, NUMBER 1 Mp5)

maxilla 1 palp uniarticulate; maxilla 2 outer plate not geniculate or gaping;
maxilliped palp article 2 not produced; pereopod 5 article 2 slender; per-
eopods 6 and 7, article 2 broad; telson entire.

Parandania boecki (Stebbing 1888)
Ie, LIS

Andania boecki Stebbing, 1888:735, pl. 36.
Parandania boecki.—Stebbing, 1899:206; 1906:95, figs. 19, 20.

Material.—Eltanin Cruise 7, Sta. 466, 12 Feb. 1963, 55°02—03’S, 44°27—
38’W, 3348-3596 m, | od.

Diagnosis.—As for genus.

Distribution.—Cosmopolitan, bathypelagic.

Remarks.—We have figured antenna 2 from our specimen since it differs
from that illustrated by Stebbing (1888, pl. 36) and J. L. Barnard (1961, fig.
27). Stebbing showed peduncle article 5 to be 3 times the length of article
4 whereas Barnard indicated article 5 to be less than twice the length of
article 4. In both cases the flagellum was longer than the peduncle. In our
specimen peduncle article 5 was 22 times the length of article 4.

Acknowledgments

The material for this study was provided by the Smithsonian Oceano-
graphic Sorting Center, Washington, D.C., and was collected under the
U.S. Antarctic Research Program. Additional specimens were generously
provided by Drs. Robert Y. George (University of North Carolina, Wil-
mington) and L. R. McKinney (Moody Marine Lab. of Texas A&M Uni-
versity). We would like to thank the following persons for their loans of
museum specimens: Dr. Denise Bellan-Santini, Station Marine d’ Endoume;
Ms. Joan Ellis, British Museum (Natural History); Ms. Elizabeth Louw,
South African Museum; Dr. H.-E. Gruner, Zoologisches Museum der Hum-
boldt-Universitat zu Berlin; Dr. Roy Olerod, Swedish Museum of Natural
History; Dr. J. K. Lowry, The Australian Museum; Prof. Jacques Forest,
Muséum National d’ Histoire Naturelle, Paris; and Dr. M. E. Christiansen,
Zoologisk Museum, Norway. This research was supported by the Smith-
sonian Oceanographic Sorting Center’s program ‘‘Cooperative Systematics
and Analyses of Polar Biological Materials’ (National Science Foundation
Grant, DPP 76-23979, B. J. Landrum, Principal Investigator). We would like
also to acknowledge the work of Ms. Patrice Rossi who produced the fig-
ures.

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Department of Oceanography, Ira C. Darling Center, University of
Maine, Walpole, Maine 04573.

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 228-239

A NEW GENUS AND SPECIES OF SPIONIDAE
(ANNELIDA: POLYCHAETA) FROM THE
NORTH AND SOUTH ATLANTIC

Nancy J. Maciolek

Abstract. —A new genus and species of spionid polychaete has been dis-
covered in collections from widespread areas in the north and south Atlantic
Ocean. Aurospio dibranchiata n. gen., n. sp., described herein, occurs from
slope to abyssal depths ranging from 300 m to 3600 m, and from areas as
far north as the Rockall Trough off Ireland to as far south as the Argentine
Basin. Aurospio 1s closely related to Prionospio, with which it is compared.
The holotype of Anaspio boreus Chamberlin, 1920 was re-examined, and
determined to be a damaged Prionospio sp., thus invalidating the genus
Anaspio.

Extensive collections of deep-sea infauna from several areas in the north
and south Atlantic have been made available for study through the work of
Drs. Howard L. Sanders and J. Frederick Grassle of the Woods Hole
Oceanographic Institution. Examination of the spionid polychaetes in these
collections has revealed a new genus and species to be a common compo-
nent of the fauna. Some specimens were found in samples previously ex-
amined by Hartman (1965), and Hartman and Fauchald (1971) and deposited
in the Allan Hancock Foundation. Additional material was obtained from
collections made by Dr. John Gage, Dunstaffnage Marine Research Labo-
ratory, in the Rockall Trough west of Ireland.

These specimens agreed with the generic diagnosis of Anaspio Chamber-
lin, 1920, which has not been reported since its original description from
Alaska. The holotype of Anaspio boreus Chamberlin was located at the
Museum of Comparative Zoology, Cambridge, Massachusetts (MCZ Cata-
log No. 2323) and re-examined. It was found to be a damaged specimen of
Prionospio sp. which had lost the first gill pair and any gills subsequent to
the third pair. (These branchial pairs are apparently lost very easily; this
condition is routinely seen in deep-sea specimens of Prionospio.) The hood-
ed hooks, which were described by Chamberlin as lacking an apical denticle,
with the process ‘‘cleft somewhat like the beak of a bird’’ (Chamberlin,
1920, p. 19B), were seen under oil immersion to be multidentate, with 6-8
teeth above the main fang. Thus, the genus Anaspio Chamberlin is consid-
ered a junior synonym of Prionospio, and the specimens reported upon
herein are referred to a new genus, Auwrospio, which is described below.

VOLUME 94, NUMBER 1 229

The holotype and a set of paratypes have been deposited in the Smith-
sonian Institution (USNM), Washington, D.C. Paratype materials have been
deposited in the following museums: Allan Hancock Foundation (AHF),
California Academy of Sciences (CAS), Museo Argentino de Ciencias Natu-
rales, Buenos Aires (MACN), British Museum of Natural History (BMNH),
and the Zoological Museum of Hamburg (ZMH). Some material has been
retained for histological and SEM examination, and some material has been
returned to the collectors: Drs. Howard L. Sanders (HLS), J. Frederick
Grassle (JFG), and John Gage (JG).

Aurospio new genus

Type-species.—Aurospio dibranchiata n. sp. Gender, feminine.

Diagnosis.—Prostomium broadly rounded anteriorly, prolonged poste-
riorly as a keel, eyes 0-2 pairs, no occipital tentacle. Peristomium partly
fused to setiger 1, not developed into wings or hood. Branchiae 2 pairs, on
setigers 3—4, cirriform, partly fused to notopodial lamellae. No interramal
or interparapodial pouches. Anterior setae capillaries, notopodial and neu-
ropodial multidentate hooded hooks posteriorly, lacking secondary hood,
one ventral sabre seta posteriorly. Pygidium with 1 long medial and 2 short
lateral cirri.

Remarks.—Aurospio is closely related to Prionospio Malmgren, 1867, in
the nature of the development of the peristomium and setiger 1, form of the
dorsal lamellae, and nature of the pygidium. The major character separating
the two genera is the initial appearance of the branchiae on setiger 2 in
Prionospio and on setiger 3 in Aurospio. As mentioned above, specimens
of Prionospio often lose the first branchial pair and those subsequent to the
third pair, and workers must be careful not to confuse damaged Prionospio
with Aurospio or other genera. However, other, more subtle differences in
the branchiae also distinguish Aurospio from Prionospio: in Aurospio the
gills appear thin and flat (in light microscopy, but see Fig. 3), and are par-
tially but obviously fused to the notopodial lamellae, while in Prionospio
the gills are stouter, rounder, and clearly separated from the dorsal lamellae.
In over 2000 specimens of Aurospio examined, 2 individuals were found
having an additional branchial pair on setiger 2, these being about as long
as the pair on setiger 3. Because the 2 specimens agreed in all other char-
acters with Aurospio, they are considered to represent anomalous individ-
uals of the genus.

Hooded hooks are present in both rami of the 2 genera, but differ in that
in Prionospio a secondary hood is present, although inconspicuous; while
in Aurospio a secondary hood is lacking.

In a recent study of Australian Spionidae, Blake and Kudenov (1978)
recognized a complex of 4 closely related genera: Paraprionospio Caullery,

230 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Streblospio Webster, Orthoprionospio Blake and Kudenov, and Prionospio
Malmgren (sensu lato), with the latter including 3 subgenera: Prionospio
(sensu stricto), Aquilaspio Foster, and Minuspio Foster. This arrangement
reflects Blake and Kudenov’s position that branchial form alone is not a
sufficient character on which generic distinctions can be based, as was sug-
gested by Foster (1971a) in her revision. In the context provided by Blake
and Kudenov, Aurospio should be considered the fifth genus in this com-
plex. A single species is known.

Etymology.—Aurum, L.—gold; spio, L.—sea nymph. The deep golden
color of the pigmented inclusions in the first setiger of many specimens
suggested the generic name.

Aurospio dibranchiata new species
Figs. 1-3

Prionospio cirrifera: Hartman, 1965 (in part; SL4, Bel, Be4, E3).—Hartman
and Fauchald, 1971 (in part; Chain cruise 58, Sta. 105). (Not Wirén, 1883).

Prionospio steenstrupi: Hartman, 1965 (in part; SL3). (Not Malmgren,
1867).

Laonice antarcticae: Hartman and Fauchald, 1971 (in part, Chain cruise
50, Sta. 87). (Not Hartman, 1953).

Material examined.—Atlantic Ocean: Gay Head—Bermuda Transect.—
Panulirus, Bermuda Sta. 1, 13 April 1960, 32°16.5’N, 64°42.5’W, 1000 m,
anchor dredge, 31 specimens (AHF). Bermuda Sta. 4, 2 May 1960,
32°17.0'N, 64°35’W, 1700 m, anchor dredge, | specimen (AHF).—Atlantis
cruise 264, Sta. HH3, 21 May 1961, 38°47'N, 70°08’W, 2900 m, anchor
dredge, 2 specimens (AHF). Sta. E3, 25 May 1961, 39°50.5’N, 70°35’ W, 824
m, anchor dredge, 20 specimens (6 for SEM, 14 AHF).—Aztlantis cruise 283,
Sta. Slope 3, 28 August 1962, 39°58.4'’N, 70°40.3’W, 300 m, anchor dredge,
1 specimen (AHF). Sta. Slope 4, 30 August 1962, 39°56.5'N, 70°39.9’W, 400
m, anchor dredge, 2 specimens (AHF).—Adtlantis IT cruise 12, Sta. 71, 24
August 1964, 38°08’N, 71°47.5'W, 2946 m, anchor dredge, 15 specimens
(AHF).—Chain cruise 50, Sta. 87, 6 July 1965, 39°48.7'N, 70°40.8’W, 1102
m, epibenthic sled, 19 specimens (AHF).—Chain cruise 58, Sta. 105, 5 May
1966, 39°56.6’N, 71°03.6’W, 530 m, epibenthic sled, 4 specimens (AHF).—
Oceanus cruise 10, Sta. 350, 10 July 1976, 38°17.0'N, 69°37.8'W, 3600 m,
box core, 11 specimens (JFG). Sta. 352, 11 July 1976, 38°16.5'N, 69°38.5'W,
3600 m, box core, 14 specimens (JFG). Sta. 367, 19 July 1976, 39°45.5’N,
70°37.2'W, 1764 m, box core, 65 specimens (JFG).—Alvin Dive, WHOI
Deep Ocean Station No. 1, 39°46’N, 70°40’W, 1760 m. Dive #407, 12 June
1972, 3 specimens (JFG). Dive #408, 14 June 1972, 2 specimens (JFG). Dive
#459, 26 September 1972, 5 specimens (JFG). Dive #460, 11 October 1972,
10 specimens (JFG). Dive #546, 5 October 1974, 1 specimen (JFG). Dive

VOLUME 94, NUMBER 1 231

#597, 30 August 1975, 4 specimens (JFG). Dive #773, 29 July 1977, 3 spec-
imens (JFG). Dive #744, 30 July 1977, 1 specimen (JFG). Dive #776, 1
August 1977, 4 specimens (JFG).—Alvin Dive, WHOI Deep Ocean Station
No. 2, 38°18’N, 65°35’W, 3600 m. Dive #657, 10 June 1976, 2 specimens
(JFG). Dive #777, 3 August 1977, 3 specimens (JFG).—Alvin Dive, Tongue
of the Ocean, 24°53.4’N, 77°40.2'W, 2032 m. Dive #753, 5 May 1977, 1
specimen (JFG). Dive #755, 12 May 1977, 2 specimens (JFG).—Ireland—
Woods Hole Transect. Chain cruise 106. Sta. 313, 17 August 1972,
51°32.2'’N, 12°35.9'W, 1491-1500 m, epibenthic sled, 586 specimens (50
specimens, HLS; 536 specimens, USNM 60343). Sta. 323, 21 August 1972,
50°08.3'’N, 13°53.7'W, 3338-3356 m, epibenthic sled, 193 specimens (USNM
60344).—Rockall Trough. Challenger cruise 12B/75, 7 September 1975, Sta.
46, 55°03.7'N, 12°06’W, 2875 m, box core, 22 specimens (JG). Sta. 47,
55°03.5’N, 12°03.5'W, 2875 m, box core, 28 specimens (JG). Sta. 48,
55°03.9'N, 12°03.9'W, 2875 m, box core, 21 specimens (JG). Sta. 49,
55°03.4’N, 12°05.3'W, 2875 m, box core, 22 specimens (JG). Sta. 50,
DRO Aa leIN eel 02,O4Wi 287 5e mm. box core, Zi specimens OG). Sta. Sik
55°03.3’N, 12°02.7'W, 2875 m, box core, 5 specimens (JG).—Challenger
cruise 9/76, 23 June 1976, Sta. 58, 54°41’N, 12°17’W, 2900 m, box core, 24
specimens (JG). Sta. 60, 56°35’N, 11°03’W, ca 2500 m, box core, 2 speci-
mens (JG). Sta. 61, 57°08’N, 12°09’W, ca 1200 m, box core, 8 specimens
(JG). Sta. 63, 56°37’N, 10°12'W, ca 1800 m, box core, 2 specimens (JG).
Sta. 65, 56°39’N, 09°40'’W, ca 1600 m, box core, 38 specimens (JG).—Chal-
lenger cruise 10/76, 1 July 1976, Sta. 68, 58°42’N, 09°43’W, ca 1800 m, box
core, 38 specimens (JG).—Bay of Biscay. Sarsia Sta. 33, 13 July 1967,
43°40.8’N, 03°36’ W, 1784 m, anchor dredge, 18 specimens (USNM 60329).
Sta. 44, 16 July 1967, 43°40.8’N, 03°35.2’W, epibenthic sled, 1739 m, 1
specimen (NJM). Sta. 65, 25 July 1967, 46°15.0'N, 04°50’W, epibenthic sled,
1922 m, 152 specimens (USNM 60330).—Canary Islands. Discovery Sta.
6697, 15 March 1968, 27°57'N, 13°46’W, 1564 m, epibenthic sled, 44 speci-
mens (USNM 60331). Sta. 6709, 18 March 1968, 27°29.8’N, 15°20’W, 2351
m, epibenthic sled, 76 specimens (USNM 60332). Sta. 6710, 19 March 1968,
27°23.6'N, 15°39.6'W, 2670 m, epibenthic sled, 71 specimens (USNM
60333). Sta. 6711, 20 March 1968, 27°13’N, 15°41’W, 2988 m, 38 specimens
(USNM 60334).—Walvis Bay—Luanda Transect. Atlantis IT cruise 42. Sta.
193, 17 May 1968, 22°56’S, 12°18’E, 2094-2191 m, anchor dredge, 1 speci-
men (USNM 60335). Sta. 200, 22 May 1968, 09°41’S, 10°55’E to 09°43.5'S,
10°57'E, 2644-2754 m, epibenthic sled, 4 specimens (USNM 60336). Sta.
201, 23 May 1968, 09°29’S, 11°34’E to 09°25’S, 11°35’E, 1964-2031 m, epi-
benthic sled, 59 specimens (USNM 60337). Sta. 202, 23 May 1968, 09°05’S,
12°17’E to 08°56'S, 12°15'E, 1427-1643 m, epibenthic sled, 47 specimens
(USNM 60338).—Argentine Basin. Atlantis IT cruise 60. Sta. 239, 11 March
1971, 36°49.0'S, 53°15.4'W, 1661-1679 m, epibenthic sled, 92 specimens (12

232 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

removed for histology, 30 paratypes USNM 60328, 10 paratypes AHF Poly
1289, 10 paratypes CASIZ type-series 1133, 10 paratypes MACN 29.610, 10
paratypes BMNH ZB1980:82-91, 10 paratypes ZMH P-16407). Sta. 264A,
28 March 1971, 36°12.7’'S, 52°42.7'W, 2041-2048 m, epibenthic sled, 52 spec-
imens (1 entire specimen, holotype USNM 60326, 51 paratypes USNM
60327).—Surinam. Knorr cruise 25. Sta. 293, 27 February 1972, 08°58.0’N,
53°04.3’W, 1456-1518 m, epibenthic sled, 169 specimens (USNM 60339).
Sta. 295, 28 February 1972, 08°04.2'’N, 54°21.3’'W, 1000—1022 m, epibenthic
Sled, 260 specimens (1 entire specimen, USNM 60340, 259 specimens
USNM 60341). Sta. 301, 29 February 1972, 08°12.4’N, 55°50.2’W, 2487-—
2500 m, epibenthic sled, 11 specimens (USNM 60342).

Diagnosis. —Prostomium broadly rounded anteriorly, extending as keel
to end of setiger 1, 0—2 pairs of indistinct red eyespots; no occipital tentacle.
Peristomium partly fused to setiger 1, not developed into wings or hood;
surrounding prostomium posteriorly as a yoke. Branchiae 2 pairs, on seti-
gers 3-4, each partially fused to dorsal lamellae, first pair longer than sec-
ond, both usually shorter than lamellae. Setiger 1 reduced; subsequent no-
topodial lamellae larger, foliaceous, becoming prolonged medially over
dorsum, largest on setigers 2—6, thereafter smaller, rounded; neuropodial
lamellae largest on setiger 3, thereafter small, rounded, displaced dorsally
in anterior setigers, ventrally placed by setiger 10. Setae of 3 types: narrow,
striated capillaries, multidentate hooded hooks, heavily granulated ventral
sabre setae. Anterior setae all capillaries, arranged in 2 tiers; neuropodial
hooded hooks from setiger 9-11 (usually 10); long-shafted notopodial hooded
hooks from setiger 24—38; ventral sabre setae from setiger 9-11. Pygidium
with | long, thin medial cirrus and 2 short, stout lateral cirri.

Description.—This is a small species, measuring up to 0.6 mm wide and
5.0 mm long for 38 setigers. The color in alcohol is white to pale yellow;
some specimens have dark golden inclusions on the dorsal edge of setiger
1, and sometimes also on the posterior tip of the prostomium.

The prostomium is broadly rounded anteriorly, extending posteriorly as
a narrow keel to the end of setiger 1 (Fig. 1A). On some specimens, a pair
of small red eyes, each composed of a cluster of pigment spots, are present
about halfway down the length of the prostomium. On a few specimens, a
second pair of eyespots is present anterior to the first pair. There is no
occipital tentacle. Some variation in the shape of the prostomium is seen;
most specimens have a prostomium as in Fig. 1A, but some prostomia were
found with tiny peaks either medially or at the lateral edges (Fig. 1B, C).
These variations are thought to be fixation artifacts, or due to the eversion
or non-eversion of the pharynx, and are not considered to be taxonomically
significant.

The peristomium is partly fused to setiger 1, appearing distinct ventrally
and laterally, but fused dorsally (Fig. 1A). The peristomium forms a yoke

VOLUME 94, NUMBER 1 233

i

Tu i
aN AY
\ tid

iM
uh Atl dy

SN
Ne Wd
" ie

Fig. 1. Aurospio dibranchiata: A, Anterior end in dorsal view; B—C, Prostomia, showing
medial and lateral peaks of some specimens; D, Everted pharynx in lateral view [Pr =
prostomium]; E, Pygidium.

around the posterior tip of the prostomium. This yoke is most obvious in
specimens which have large amounts of the golden pigment mentioned
above. A pair of grooved palps are present at the junction of the peristomium
and prostomium, and extend about one-third to one-half the length of the
body. The palps were lost in all but a few (3-4) of the specimens examined.

The pharynx is an eversible pouch, heavily muscularized on the ventral
surface, and ciliated on the anterodorsal surface (Fig. 1A, D).

Branchiae are present on setigers 3—4. They are short and fused for one-

234 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Aurospio dibranchiata: A, Notopodia and branchiae of setigers 3 and 4 in posterior
view; B, 15th setiger, with enlargement of sabre seta, in anterior view; C, 30th setiger in
anterior view; D, Neuropodial hooded hook, inset [not to scale] showing arrangement of teeth.

quarter to one-third their length to the dorsal lamellae, beyond which they
do not extend, and by which they are usually completely hidden (Figs.
1A, 2A, 3). The gills on setiger 3 are longer than those on setiger 4,
which are stubby in appearance. Cilia line the medial edge of the gills on
setiger 3, and continue across the dorsum (Fig. 3A, B), but are not present
on the gills on setiger 4.

The notopodium of setiger 1 is reduced; on setigers 2—6, the notopodial

VOLUME 94, NUMBER 1

N
ios)
(Nn

A

'*.

: . pal

Fig. 3. Aurospio dibranchiata: A, SEM of anterior end in dorsolateral view [scale = 20
ym]; B, SEM of branchiae on setigers 3 and 4 [scale = 4 um].

236 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

lamellae are large, broad and foliaceous, with the medial edge prolonged
over the dorsal surface. From setigers 6—14 the lamellae become increas-
ingly smaller. A few specimens appear to have very low dorsal ridges on
setigers 5—10, but the appearance of this character is variable within the
species.

The neuropodium of setiger | is also reduced to a small, rounded lamella,
and displaced dorsally. On setiger 2 it is a small auricular lobe which be-
comes larger on setiger 3 and then smaller on subsequent setigers. The
neuropodia of setigers 2-10 are increasingly ventral in position (Fig. 1A).
In posterior setigers, both noto- and neuropodia are small, rounded lobes
(Fig. 2B, C).

Setae consist of 3 types: capillaries, multidentate hooded hooks and ven-
tral sabre setae. Anterior setae are all capillaries, arranged in 2 tiers. These
setae are often extremely long, particularly the neurosetae in setiger 2 (Fig.
1A), but also both the noto- and neurosetae posteriorly. They appear nar-
row, uni- or bilimbate (depending on the angle at which they are viewed;
see Foster, 1971b), and striated with punctations along the striae. Neuro-
podial hooded hooks appear from setiger 10, occasionally from setiger 11,
and less frequently from setiger 9. Long-shafted notopodial hooks appear in
the last few setigers (24-38), and will probably not be seen unless complete
specimens are collected. These hooks appear quadridentate if viewed from
the side (Fig. 2D), but actually have 3 pairs of teeth arranged above the
main denticle (Fig. 2D—inset). There is no secondary hood. Hooks number
6-9 per neuropodial ramus, and 2-3 per notopodial ramus. Accompanying
capillaries number up to 8 in the neuropodium; up to 3 in the notopodium.
A single heavily granulated and bilimbate ventral sabre seta is present from
setiger 10 (9-11) (Fig. 2B). Rarely, a second sabre seta is present.

The pygidium is rarely retained in entirety. The pygidium of the holotype
has | long medial cirrus and 2 short, stout lateral cirri (Fig. 1E).

Remarks.—Aurospio dibranchiata is an extremely widespread species,
occurring over most of the Atlantic Ocean (Fig. 4), and from slope depths
of 300 m to abyssal depths of 3600 m. Twenty-six of the 30 records are from
depths greater than 1000 m. The records from shallower depths are all from
the Gay Head—Bermuda transect, although stations in less than 1000 m were
sampled on most of the other transects.

A. dibranchiata was the second most common species, comprising 5.1
percent of the fauna, in core samples taken by Grassle (1977) at 1760 m off
Woods Hole. It was also found in samples taken from recolonization ex-
periments conducted at the same site. Almost nothing is known about its
life history: only one specimen containing oocytes was found; these mea-
sured 50-90 microns for the greatest visible dimension.

The consistency of taxonomic characters in A. dibranchiata over its depth
and geographic range is notable. Neuropodial hooded hooks began in setiger

VOLUME 94, NUMBER 1 2By)

LIC 100, 90 80 70) OO) HO YOr sO 5200 Io)
b Rockall
Trough

Ireland- ~—
Woods Hole
, Bay o&
Biscay
Gay Head- °
Bermuda

a
o, e
e

Canary Se

hee

Walvis Bay-
Luanda

Agraeotine Basin

Fig. 4. Sampling transects in the north and south Atlantic Ocean from which Aurospio
dibranchiata was collected.

238 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

10 in nearly every specimen examined. Only 2 or 3 individuals were found
in which hooded hooks began in setiger 9. In the 4 samples from the Canary
Islands, roughly 10 to 25 percent of the specimens had hooded hooks starting
in setiger 11, suggesting a small variation on a regional basis. Other char-
acters, such as shape of prostomium, length of setae in setiger 2, shape of
parapodial lamellae and presence or absence of dorsal crests varied little
between or among samples.

The golden pigment in the peristomium/first setiger region is most pro-
nounced in the material from the Argentine Basin, and present although
somewhat less obvious in most specimens from Surinam and many speci-
mens from other locations. Initial histochemical tests reveal this material to
be a lipofuscin. Further histological work is in progress on this and other
spionid species in which a similarly pigmented material is found (Maciolek,
Hillman and Lahey, in prep.).

Acknowledgments

I am grateful to Drs. H. L. Sanders, J. F. Grassle and J. Gage for pro-
viding the material for this study, to Dr. J. A. Blake for the SEM photo-
graphs, and to California State University, Fresno, for use of the Zeiss
Novascan 30 SEM. Curators at the Allan Hancock Foundation and the
Museum of Comparative Zoology were very helpful in meeting requests for
material. Drs. J. A. Blake and K. Fauchald were generous in offering critical
remarks and guidance. Mr. G. Hampson and Mrs. L. Morse-Porteous
helped in assembling station data, and Mrs. I. Williams commented on the
manuscript. This work represents a portion of a thesis to be submitted to
Boston University in partial fulfillment of the requirements for the degree
of Doctor of Philosophy, and was supported in part by a Sigma Xi Grant-
in-Aid-of-Research. The study was carried out partly at the Marine Biolog-
ical Laboratory, Woods Hole, and partly at Battelle’s William F. Clapp
Laboratories, Duxbury, Massachusetts.

Literature Cited

Blake, J. A., and J. Kudenov. 1978. The Spionidae (Polychaeta) from southeastern Australia
and adjacent areas with a revision of the genera—Mem. Nat. Mus. Victoria 39:171-—
280.

Chamberlin, R. V. 1920. Polychaeta.—Report Canadian Arctic Exped. 1913-1918, 9B:1-41B.

Foster, N. M. 197la. Spionidae (Polychaeta) of the Gulf of Mexico and the Caribbean Sea.—

Stud. Faun. Curagao Caribb. Islands 36:183 pp.

. 1971b. Notes on spionid (Polychaeta) setae.—Trans. Amer. Microscop. Soc. 90(1):

34-42.

Grassle, J. F. 1977. Slow recolonisation of deep-sea sediment.—Nature 265(5595):618-619.

Hartman, O. 1965. Deep-water benthic polychaetous annelids off New England to Bermuda
and other North Atlantic areas.—Allan Hancock Found. Occ. Pap. 28:378 pp.

VOLUME 94, NUMBER | e339)

, and K. Fauchald. 1971. Deep-water benthic polychaetous annelids off New England
to Bermuda and other North Atlantic areas. Part 2.—Allan Hancock Monogr. Mar. Biol.
6:327 pp.

Battelle’s New England Marine Facility, The William F. Clapp Laborator-
ies, Inc., P.O. Drawer AH, Duxbury, Massachusetts 02332.

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 240-253

RECLASSIFICATION AND REDESCRIPTION OF THE
COMATULID COMATONIA CRISTATA (HARTLAUB)
(ECHINODERMATA: CRINOIDEA)

Charles Garrett Messing

Abstract. —Comatonia cristata (Hartlaub) is redescribed and illustrated
on the basis of new and existing material. Well-developed pinnular combs,
the only character relating this species to the Comasteridae, are modifica-
tions of rudimentary combs found in heliometrine antedonids. Comatonia
agrees with all diagnostic characteristics of, and is here transferred to, the
family Antedonidae, approaching most closely the heliometrine Anthometra
adriani (Bell). Cirrus morphology may be convergent with sympatric Coc-
cometra hagenii (Pourtales).

The comatulid genus Comatonia A. H. Clark contains a single species,
C. cristata (Hartlaub), found primarily at moderate depths off the south-
eastern United States. A. H. Clark (1916, 1921, 1931) included it in the
family Comasteridae (suborder Comasteracea) entirely on the basis of the
well-developed combs on the first pair of oral pinnules. Such combs, the
teeth of which are flat, usually triangular or spade-shaped expansions of the
aboral surface of distal pinnule segments, are diagnostic of this family. In
all other respects, however, Comatonia belongs in the family Antedonidae
(suborder Antedonacea). Although Gislén (1924) proposed that the species
be referred to this family, he was not followed by later authors (Clark, 1931;
Messing, 1975; Rasmussen, 1978) with the exception of Meyer (1972). In
the course of revising the extant crinoids of the tropical western Atlantic,
I have discovered that the comb teeth of Comatonia appear to be modifi-
cations of comb rudiments found in some antedonid genera; they are con-
vergent with rather than homologous to comasterid comb teeth. No reason
remains for retaining this comatulid within the Comasteridae. Its placement
within the Antedonidae is discussed below.

In addition, new material, collected primarily by the University of
Miami’s R/V Gerda, together with a re-examination of existing collections,
has revealed several previously unrecorded variations. I have, therefore,
redescribed and illustrated the species as follows. The tabular arrangement
of data follows that of A. M. Clark (1970) and Messing (1978). USNM,
UMML and MCZ refer to National Museum of Natural History, University
of Miami Marine Laboratory (Rosenstiel School of Marine & Atmospheric
Science) and Museum of Comparative Zoology (Harvard University) cata-
logs, respectively.

VOLUME 94, NUMBER 1 241

Genus Comatonia A. H. Clark (1916)

Diagnosis.—Radials 5; arms 10, aborally rounded; cirri slender and nu-
merous, up to 20 mm with 23 cirrals and lacking aboral processes; comb on
P, and P.,, rarely absent, rarely present on P,, occupying all but proximal
third of pinnule and composed of large, round teeth; one or more proximal
pinnulars of P, sometimes longer than wide; Br.—Br, (sometimes Br,—Br,,)
each with spinose, median, aboral knob or ridge; division series lacking
spines. (Modified from A. H. Clark, 1931:289.) Type-species: Actinometra
cristata (Carpenter MS.) Hartlaub, 1912, by original designation and mono-
typy (A. H. Clark, 1916:115).

Comatonia cristata (Hartlaub)
INES, N—3, 4D, ©, IMs ly SA

Antedon sp. Carpenter, 1881:155.

Actinometra cristata (Carpenter MS.) Hartlaub, 1912:280, 413, 473-475; pl.
10, figs. 1-5; pl. 15, figs. 10-11.

Comatonia cristata: A. H. Clark, 1916:115.—H. L. Clark, 1918:7, 9.—A.
ee Clarke OD E144 24232 2S 284 292293), 805, 372) 595, 619:
1931:57, 61, 63, 64, 78, 82, 87, 88, 89, 231, 288-292, 399, 400; pl. 6.—
Meyer, 1972:64.—Meyer et al., 1978:416—417, 428, 430, 431, 432.

Description.—Centrodorsal hemispherical or low, rounded conical, 1.3-
4.4 mm across; DH 1.3-1.8 (Figs. la, d, e). Cirrus sockets crowded, ar-
ranged in spiral whorls overlain by irregular columns of 3-6 sockets and
almost completely covering centrodorsal. Aboral pole small, not more than
0.4 x basal diameter of centrodorsal, flat or convex, almost smooth to
strongly papillose, rarely depressed and ringed with papillae; when present,
papillae arising, one each, from obsolete, apical sockets. Centrodorsal mar-
gin midradially concave; interradial projections rounded or triangular; cen-
tral cavity broad and deep, with prominent oral lip and slightly more than
half basal diameter of centrodorsal; interior, lateral walls of cavity bearing
narrow, interradial buttresses; oral opening pentagonal or irregularly scal-
loped (Fig. 2)).

Cirri (Figs. 2a—g) slender, about L-C, 12—23, up to 20 mm long, laterally
compressed distally. Apical cirri shorter, weaker and composed of fewer
cirrals than peripheral cirri. First cirral short; second squarish; third with
LW almost 2.0; third to sixth (usually fifth) longest, with LW 2.3—4.0; fol-
lowing cirrals decreasing in length but remaining longer than wide; ante-
penultimate squarish; all but basal few cirrals somewhat expanded distally;
aboral spines or carinae absent; opposing spine directed obliquely distally;
terminal claw curved and slightly shorter than penultimate cirral.

Rosette at or just below aboral surface of radial pentagon, bearing large,
orally curved, flaring, radial processes and short, triangular, interradial pro-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

we)
aN
Ww

Fig. 1. Comatonia cristata: a, Radial side view of a specimen (USNM E17863) showing
centrodorsal, bases of five cirri and proximal part of three rays (partly shaded); b, Radial
articular facet (USNM E19971) showing calcareous deposit bridging high, intermuscular walls;
c, Another radial facet from same specimen; d, Interradial side view of centrodorsal (USNM
E19971); e, Same (USNM E19188); f, Proximal part of single postradial series (IBr, to Brs),
aboral view (USNM E19971); g, Oral view of rosette (USNM E19188); h, Same, radial side
view. Scale: a-f = 2 mm; g—h = | mm.

VOLUME 94, NUMBER 1 243

Fig. 2. Comatonia cristata: a, Peripheral cirrus (USNM E19971); b, Apical cirrus, same
specimen; c, Peripheral cirrus (USNM E19177); d, Cirrus of small specimen (USNM E19971);
e, Peripheral cirrus (USNM E17863); f, Apical cirrus, same specimen; g, Apical cirrus, same
specimen as c; h, Proximal brachials (Br,—Br,,), lateral view, showing bases of P,, P,, P, and
spinose, aboral knobs on Br,—Br, (USNM E19177); i, Radial side view of small specimen
(USNM E19971) showing centrodorsal, bases of several cirri and proximal part of 3 rays; j,
Centrodorsal, oral view, with lip cut away at left to show actual extent of central cavity (USNM
E19188). Scale: a-g = 4 mm; h-j = 2 mm.

244 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

cesses (Figs. lg, h). Basal rays absent; possible vestiges in one specimen
fused to centrodorsal (Fig. 2)).

Radials visible only interradially or as extremely narrow band (Fig. 1a).
Articular facet slightly longer along oral-aboral axis than wide, deeply ex-
cavated and not parallel to oral-aboral axis of specimen; muscular fossae
about twice height of interarticular ligament fossae, thin and laminar, flat or
overhanging orally, separated by low, narrow ridge. In a single specimen
(USNM E19971, Figs. 1b, c), a pair of thin, irregular septa project distally
(along the longitudinal axis of the ray) from the margins of the intermuscular
ridge to the level of the central canal opening; these vary among the facets
and, at one point on one facet, fuse together, creating the illusion of a double
central canal (Fig. 1b).

Division series (Figs. la, f) separated. IBr, very short, usually with me-
dian tubercle and concave distally. Primaxil (IBr.) rhombic or triangular,
round and swollen proximally, forming, with median tubercle of IBr, , prom-
inent synarthrial swelling; distal half flat, narrow and acute; WL 1.2-1.3.

Arms increasing slightly in width from base to Br.—Br, before tapering;
longest attached fragment 40 mm; estimated original length up to 70 mm.
Br, very short, longer exteriorly, united interiorly over primaxil, deeply
incised distally by prominent, usually swollen, proximal synarthrial projec-
tion of large, irregularly quadrate Br, (Figs. la, f). Br;,, oblong, up to 2.1
mm across; WL 1.2-1.7. Br; to Br, oblong or slightly trapezoidal, rarely
bearing prominent, alternating articular swellings; WL 1.7—2.5. Bry,,,. ob-
long; following brachials trapezoidal, becoming triangular between Br,, and
about Br,;; WL 1.3-2.0.

Spinose, median, aboral knob or longitudinal ridge present on Br.—Br,,
sometimes present but weak as far as Br,,, disappearing abruptly thereafter
or incorporating into finely crenulate distal margin of brachials, usually ab-
sent from syzygial pairs but sometimes present on Br, (Fig. 2h). In smallest
individual (Fig. 21), synarthrial swellings reduced, brachials elongated after
Br,; and never triangular.

Syzygies at Br,,,, Bro,,,, usually Br,,,,,; (sometimes Br,;,,,); subsequent
interval three or, rarely, four.

P, (Figs. 3c, j) flagelliform, up to 14 mm with as many as 52 pinnulars; all

=>

Fig. 3. Comatonia cristata: a, P,, lacking comb (USNM E19971); b, P;,, same specimen and
arm; c, P, (USNM E19188); d, P,, same specimen and arm; e, P,, with comb developed on both
sides of pinnule; arrow denotes pinnular with comb tooth similar to that of Anthometra adriani
(USNM E19971); f, P, (USNM E19177); g, P,,, same specimen and arm; h, P,., same; i, P,,
same; j, P,, small specimen (USNM E19971); k, P, with rudimentary comb (USNM E17863).
Scale: 2 mm.

245

VOLUME 94, NUMBER 1

246 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

but proximal 8-12 pinnulars each bearing a flat, round, somewhat distally-
directed comb tooth at least as high as width of pinnular bearing it and
sometimes 2—3x higher; proximal 3—6 pinnulars short and rhombic; subse-
quent pinnulars squarish to twice as long as wide but appearing short due
to relative height of comb teeth; teeth rarely developed on both sides of
pinnular (Fig. 3e). P, (Fig. 3d) basally stouter than P,, up to 10 mm with 18
pinnulars but often only half length of P,; proximal 2 pinnulars short; third
squarish; following of about equal length (except near tip) but with LW
increasing from about 1.3 to 4.0 as pinnule tapers; all except basal 2 with
finely spinose, expanded, distal margins or with 1 or 2 stout, distal, aboral
spines. P.,, occasionally bearing a comb similar to that of P, but with fewer,
less well-developed teeth or with relatively few, low, distal teeth on pin-
nulars similar to those of combless P, (Fig. 3k). Following pinnules similar
to combless P, but with second segment becoming abruptly tapered. P,
slightly longer or shorter than P,, of about 11 segments. P, up to 10 mm
with 18 pinnulars. Genital pinnules (Fig. 31) stouter than distal pinnules.
Longest intact distal pinnules incomplete, at least 10 mm with at least 19
segments. P,, 6.5 mm with 15 pinnulars on arm of specimen bearing P, of
5.5 mm with 11 segments. Interior pinnules similar to exterior ones (Figs.
3a, b, e-1). Single P, lacking comb and resembling subsequent pinnules
though shorter and composed of fewer segments than succeeding P,, (Figs.
3a, b).

Gonads borne on third through sixth to eighth pinnulars from P,—P, (rare-
ly, P, or P, and, perhaps in one specimen, P,) to at least P,,. Pinnule am-
bulacra lined with fine, unbranched rods sometimes slightly expanded and
perforated at one end.

Tegmen naked; mouth central or subcentral; anus submarginal; a few,
small dark saccules present on pinnule ambulacra in two specimens (USNM
E548, 34634).

Color.—In life, yellow. In alcohol, white or pale brown, sometimes with
dark brown ambulacra, gonads, and tegmen.

Biology.—A. C. Neumann and J. C. Lang, during a DSRV Alvin dive,
collected a single individual in 580 m clinging to a fragment of the branching,
ahermatypic coral Lophelia prolifera (Pallas) and in association with large
numbers of the small comasterid Comatilia iridometriformis A. H. Clark
which is usually found on this coral (Messing, unpublished). Most specimens
have been collected with the Pourtales Terrace south of the Florida Keys
apparently on hard, sediment-free substrates and usually in association with
large numbers of the antedonid Coccometra hagenii (Pourtales).

Material examined.—NORTH CAROLINA: Fish Hawk sta. 7302
(USNM 34634, 1 specimen). POURTALES TERRACE (SOUTH OF THE
FLORIDA KEYS): Gerda sta. 220 (UMML 44-169, 1), Gerda sta. 589
(USNM E19178, 1), Gerda sta. 840 (USNM E19180, 1), Gerda sta. 865

VOLUME 94, NUMBER 1 247

(USNM E19177, 1), Gerda sta. 977 [USNM E17863, 1; UMML 44-174, 2
(1 sent to D. B. Macurda, Jr.)], Gerda sta. 978 (USNM E19188, | disso-
ciated), Gerda sta. 1102 (USNM E19971, 4, 1 dissociated), Fish Hawk sta.
7298 (USNM 34628, 1), Univ. Iowa Bahama Exped. 1893, sta. 48 (USNM
E4290, 1 dissociated, not Univ. lowa Barbados-Antigua Exped. as labelled;
MCZ 747, 1), Eolis, J. B. Henderson coll. (USNM E548, 1), No expedition
data (MCZ 800, 2; removed from MCZ 755). BAHAMAS: Alvin dive 761
(USNM E19181, 1). YUCATAN CHANNEL: Gerda sta. 885 (USNM
E19179, 1).

Type.—Carpenter (1881) mentioned a new species of Antedon undoubt-
edly referable to Comatonia cristata as occurring with Antedon hagenii
(=Coccometra hagenii) in the Straits of Florida. Hartlaub (1912) described
in detail a single specimen accompanied by the label ‘‘Com. 101, Stn. —”’
and suggested that it possibly came from the Challenger rather than from
the Blake collection. At the end of his description (p. 475), he gave the
locality as unknown.

A. H. Clark re-examined the specimen and reinterpreted the label as
reading “‘Corw. Po. 1, Stn. __”’ which he identified as Corwin station 1P
(5 mi. SSW of Sand Key, Florida, 164-183 m, 17 May 1867) (1931:292). He
also wrote that Hartlaub had *‘a single specimen of this species without a
locality label,’ a lapse resulting from Hartlaub’s listing of the locality as
unknown the CORWIN specimen, according to Clark, is in the MCZ
(cat. no. 7), but I have not seen it.

Distribution.—Cape Lookout, North Carolina; Straits of Florida; Yucatan
Channel. All but 3 specimens were collected on the Pourtales Terrace south
of the Florida Keys. Possible depth range: 14-580 m. Confirmed depth
range: 14—580 m. For material collected on the Pourtales Terrace, the ranges
are 146-411 and 152-306 m, respectively. The Cape Lookout specimen was
taken in 14 m and the specimen from Arrowsmith Bank, Yucatan Channel
in 419-434 m. (Emended from Meyer et al., 1978:417.)

The specimen collected by Alvin in 580 m represents the only record from
the insular margin of the Straits. Its greater depth relative to continental
margin records parallels records for its frequent associate, Coccometra hag-
enii. The latter occurs primarily between 150 and 250 m on the Pourtales
Terrace (locality data for the 2 records in excess of 300 m here fall in less
than 250 m on recent bathymetric charts) and as deep as 442 m off Cuba.
Two records from the Blake Plateau are still deeper (805 and 1046 m) and
one of these includes Lophelia.

Discussion.—A. H. Clark (1921, 1931) considered Comatonia, Comatilia
and Comatulides as ‘‘primitive generalized’’ (1921:619) members of the fam-
ily Comasteridae. He included Comatonia in the subfamily Capillasterinae
because of its supposed resemblance to the small, 10-armed, monotypic
genera Comatilia and Microcomatula and despite the absence of diagnostic,

248 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

aboral, cirral processes. Comatonia and Comatilia both occasionally bear
saccules along their ambulacra but the resemblance ends here. Comatilia
appears to be paedomorphic; its ““generalized’’ appearance (e.g., 10 arms,
central mouth) along with other characteristics (e.g., large radials, deficient
pinnulation, elongated cirrals and brachials) are the result, I believe, of
accelerated sexual maturation and persistent juvenile, morphology (proge-
nesis sensu Gould, 1977) (Messing, unpublished). Microcomatula mortenseni
A. H. Clark is known from a single, incompletely described specimen. Al-
though discussion of its affinities must await re-examination of the type, the
published description (A. H. Clark, 1931) also agrees with specimens of the
comb-bearing antedonid Ctenantedon kinziei Meyer in some respects.

Comatonia agrees with the suborder Antedonacea and family Antedoni-
dae in all diagnostic characters (Clark and Clark, 1967; Rasmussen, 1978).
It also exhibits non-diagnostic traits, including its cirrus arrangement and
morphology and prominent synarthrial swellings (Gislén, 1924:229, foot-
note), found in many antedonids but unknown in the Comasteridae. Its
typically antedonid centrodorsal, almost covered with cirrus sockets, occurs
in the Comasteridae only in Microcomatula which, as mentioned above,
also may not belong in this family. Meyer (1972) noted that its radial articular
facets and large centrodorsal cavity are characteristically macrophreatine
(antedonacean) and agreed with Gisleén that its affinities lay with this sub-
order rather than with the comasterids.

Following Gislén (1924:229, footnote), I believe Comatonia most closely
approaches the antedonid subfamily Heliometrinae. Although the antedonid
subfamilies are not uniformly well-defined, I here follow Clark and Clark’s
(1967) treatment. Some heliometrine species bear rudimentary or moderate-
ly developed combs on their oral pinnules, although these are never nearly
as developed as in Comatonia and are weaker than in most comasterids. In
the latter, comb tooth rudiments usually develop from the middle of the
aboral side of the pinnular. In Comatonia and the heliometrines Anthometra
and Florometra, the comb teeth develop as modifications of the spinose,
distal, aboral margin of pinnulars proximal to the comb. As the teeth in-
crease in size on more distal pinnulars, they expand proximally, occupying
more and more of the aboral surface of the pinnular. In Comatonia, this
derivation is most obvious in small specimens and on comb-bearing P, (Figs.
3f, j, k). It is obscured in larger specimens in which pinnulars proximal to
the comb lack spinose, distal margins. Figures 4 and 5 show the close re-
semblance of rudimentary comb teeth and pinnular articular facets in Coma-
tonia and the heliometrine Anthometra adriani (Bell) as well as the strong
distinction between these characters in these two species and corresponding
structures in the comasterid Neocomatella alata (Pourtales). The arrow in
Fig. 3e indicates an apparently regenerating, comb tooth-bearing pinnular
in Comatonia that is similar to fully developed teeth in Anthometra adriani

249

VOLUME 94, NUMBER |

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. a, Distal articular facet of proximal pinnular, Neocomatella alata; b—c, Same, Com-
atonia cristata, stereo pair; d, Middle pinnular with rudimentary comb tooth, ambulacral view,
Anthometra adriani; e, Same, abambulacral view, N. alata; f, Distal articular facet of proximal
pinnular, A. adriani; g, Middle pinnular with rudimentary comb tooth (same pinnule but slightly
distal to d), ambulacral view, A. adriani; h-i, Same, C. cristata, stereo pair. Scales: 0.1 mm.

(Fig. 4h). In the heliometrine genus Florometra, comb teeth normally do
not develop beyond the rudimentary, distal, spinose stage.*

* A.M. Clark (personal communication) has brought to my attention the fact that another
antedonid, Annametra occidentalis (A. H. Clark) (subfamily Antedoninae), may also bear

VOLUME 94, NUMBER |

Fig. 5. a-—g, Comatonia cristata: a, Distal pinnular with well developed comb tooth, abam-
bulacral view; b—c, Same, ambulacral view, stereo pair; d, Middle pinnular with moderately
developed comb tooth, ambulacral view; e-f, Middle pinnular (more proximal than d, with less
well developed comb tooth), ambulacral view, stereo pair; g, Same, abambulacral view; h,
Anthometra adriani (Bell), distal pinnular with well developed comb tooth, ambulacral view;
i, Neocomatella alata (Pourtales), same, obliquely ambulacral view. Scales: 0.1 mm.

rudimentary comb teeth on Pl. It appears from Figure 6 in Clark and Clark (1967:95) that
these teeth are also derived from the distal aboral pinnular margin as discussed above. I
have not yet examined specimens of this species.

Day PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Within the Heliometrinae as currently construed, Comatonia appears
most similar to Anthometra adriani. Both bear a median, aboral knob or
ridge on the proximal brachials, although these are more prominent and
occur on more brachials in Anthometra. Both have only the first pair of
pinnules modified (rarely, P, in Comatonia). The radial articular facets of
the two species, and of another heliometrine, Solanometra antarctica (Car-
penter), are also closely similar. The paired, irregular “‘septa’’ projecting
distally (along the longtiudinal axis of the ray) from the sides of the inter-
muscular ridge in one specimen of Comatonia vary among the facets, appear
to represent a secondary deposition of stereom and do not alter the basic
architecture of the facet.

The relatively short, slender cirri of Comatonia differ from those of other
heliometrine species although they do resemble those of juvenile Anthomet-
ra (centrodorsal diameter 1.5 mm). They are also similar, however, to cirri
of the thysanometrine antedonid Coccometra hagenii. Comatonia and Coc-
cometra hagenii have almost identical, restricted ranges: from the Mexican
side of the Yucatan Channel to Cape Lookout, North Carolina. They occur
sympatrically and in the greatest abundance on the Pourtales Terrace (Clark
and Clark, 1967; Messing, 1975; Meyer et al., 1978). The terrace is largely
sediment-free, is subject to strong currents, exhibits some karst-like topog-
raphy (Jordan, 1954; Jordan et al., 1964; Gorsline and Milligan, 1963) and
apparently represents a distinctive, if not unique, environment in the trop-
ical western Atlantic Ocean. That Comatonia and Coccometra hagenii are
virtually the only crinoids ever collected here despite intensive dredging
suggests that their cirrus morphology may be adaptively convergent.*

Comatonia conforms least to the Heliometrinae in terms of distribution.
It occurs tropically and subtropically at moderate depths, whereas the re-
maining genera are found in polar, subpolar and cold, eastern Pacific waters.
It is morphologically distinct enough, however, to suggest that it diverged
from heliometrine stock before they became exclusively cold-water forms.
The Bathymetrinae, allied to the Heliometrinae through Anthometra ac-
cording to A. H. Clark (Clark and Clark, 1967), is a primarily deep and cold-
water subfamily that includes species that enter warmer water, e.g., Tri-
chometra cubensis (Pourtales). Despite the relatively low latitudes at which
it occurs, steep isothermal tilting across the Straits of Florida and Yucatan
Channel indicates that Comatonia most likely occurs in water of less than
10°C.

Acknowledgments

I am grateful to Drs. Frederick M. Bayer of the Department of Inverte-
brate Zoology, National Museum of Natural History and Gilbert L. Voss
of the University of Miami for making material collected by R/V Gerda

* Despite these similarities, they do not appear to be closely related. In C. hagenii, the low hemispheric centrodorsal has a
broad, smooth aboral pole; the cirri, though similar to those of Comatonia, are more delicate and are lost far more often; the arms,
usually broken near the base in Comatonia, are far more often preserved in C. hagenii. Although P1 in both species is composed
of many short pinnulars which may also be adaptively convergent, in C. hagenii, as in other thysanometrine species, the corners
of the pinnulars are cut away giving the pinnule a distinctively beaded appearance.

VOLUME 94, NUMBER | 253)

available to me and, with Drs. Lowell P. Thomas and Donald R. Moore
(both of the University of Miami), for supervising my master’s thesis re-
search, part of which is here included. I am indebted to Dr. David L. Pawson
also of the Department of Invertebrate Zoology for supervising my post-
doctoral research and, with Dr. Robert Woollacott of the Museum of Com-
parative Zoology, Harvard, for making available material from their re-
spective institutions. The scanning electron micrographs were taken by Ms.
Mary-Jacque Mann, of the National Museum of Natural History.

Literature Cited

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Harvard 9(4):151—169; 1 pl.

Clark, A. H. 1916. Seven new genera of echinoderms.—J. Wash. Acad. Sci. 6(5):115—122.

. 1921. A monograph of the existing crinoids 1(2).—Bull. U.S. Natl. Mus. 82:xxv +

TESS SI 8.

. 1921. A monograph of the existing crinoids 1(3).—Bull. U.S. Natl. Mus. 82:vii + 816;

86 pls.

, and A. M. Clark. 1967. A monograph of the existing crinoids 1(5).—Bull. U.S. Natl.

Mus. 82:xiv + 860.

Clark, A. M. 1970. Marine Invertebrates of Scandinavia, 3. Echinodermata Crinoidea.—
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Clark, H. L. 1918. Report on the Crinoidea and Echinoidea collected by the Bahama Expe-
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Gislen, T. 1924. Echinoderm Studies—Zool. Bidr. Uppsala 9:iv + 316.

Gorsline, D. S., and D. G. Milligan. 1963. Phosphatic deposits along the margin of the Pour-
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Gould, S. J. 1978. Ontogeny and Phylogeny.—Belknap Press, Cambridge, Mass. xvi +

501 pp.

Hartlaub, C. 1912. Reports on the results of dredging . . . by the U.S. Coast Survey Steamer
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491; 18 pls.

Jordan, G. F. 1954. Large sinkholes in the Straits of Florida.—Bull. Am. Assoc. Petr. Geol.
38: 1810-1817.

Jordan, G. F., R. J. Malloy, and J. W. Kofoed. 1964. Bathymetry and geology of Pourtales
Terrace, Florida.—Mar. Geol. 1(3):259-287.

Messing, C. G. 1975. The systematics and distribution of the Crinoidea Comatulida (exclusive

of the Macrophreatina) collected by the R/V Gerda in the Straits of Florida and adjacent

waters.—M.S. Thesis, Univ. of Miami, Coral Gables, Fla. 296 pp.

. 1978. A revision of the comatulid genus Comactinia A. H. Clark (Crinoidea: Echi-

nodermata).—Bull. Mar. Sci. 28:49-80.

Meyer, D. L. 1972. Ctenantedon, a new antedonid crinoid convergent with comasterids.—
Bull. Mar. Sci. 22(1):53-66.

Meyer, D. L., C. G. Messing, and D. B. Macurda, Jr. 1978. Zoogeography of tropical western
Atlantic Crinoidea.—Bull. Mar. Sci. 28:412-441.

Rasmussen, H. W. 1978. Articulata in: R. C. Moore and C. Teichert, eds., Treatise on
Invertebrate Paleonotology, Part T, Echinodermata 2(3):T813-—T937.

Department of Invertebrate Zoology, National Museum of Natural His-
tory, Smithsonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 254-263

A NEW SPECIES OF TEGASTES
(COPEPODA: HARPACTICOIDA) ASSOCIATED WITH A
SCLERACTINIAN CORAL AT ENEWETAK ATOLL

Arthur G. Humes

Abstract.—The harpacticoid copepod Tegastes acroporanus, new
species, is associated with the scleractinian coral Acropora florida (Dana)
at Enewetak Atoll. Until now only one harpacticoid species, Tegastes geor-
gei Marcus and Masry, 1970, from Stylophora and Pocillopora in the Gulf
of Elat, has been known to be associated with hard corals. The two species
of Tegastes from corals resemble each other in the form of leg 5 in the
female, but differ in the nature of the setae on the exopod of leg 4 and in
the degree of sexual dimorphism in the first antenna of the male.

Although many cyclopoid copepods are associated with Scleractinia
(Humes, 1979), harpacticoids have not developed associations with hard
corals to the same extent. At present the only harpacticoid known to live
with corals is Tegastes georgei Marcus and Masry, 1970. These authors
found it with Stylophora and Pocillophora in the Gulf of Elat.

The specimens from the Marshall Islands reported here were collected by
the author and Dr. Charles T. Krebs during field work made possible by the
support and facilities of the Enewetak Marine Biological Laboratory,
Enewetak. (The name of the atoll was formerly spelled Eniwetok.)

I thank Dr. John W. Wells, Department of Geological Sciences, Cornell
University, who very kindly identified the coral host.

The observations and measurements were made on specimens cleared in
lactic acid. All figures were drawn with the aid of a camera lucida. The
letter after the explanation of each figure refers to the scale at which it was
drawn. The abbreviations used are: R = rostrum, A, = first antenna, A, =
second antenna, and L = labrum.

Harpacticoida G. O. Sars, 1903
Tegastidae G. O. Sars, 1904
Tegastes Norman, 1903

Tegastes acroporanus, new species
Figs. 1-26

Type-material.—109 22, 85 36 from Acropora florida (Dana), in 2 m,
western end of Bogon Island, Enewetak Atoll, Marshall Islands, 23 June
1969. Holotype 2, allotype, and 155 paratypes (90 22, 65 6d) deposited

VOLUME 94, NUMBER 1 PENS)

Figs. 1-7. Tegastes acroporanus, female. 1, Lateral (A); 2, Dorsal (A); 3, Genital and
postgenital segments, lateral (B); 4, Genital segment, with outline of postgenital segments
shown by broken line, ventral (B); 5, Caudal ramus, lateral (C); 6, Egg sac, lateral (A); 7,
Rostrum and part of labrum, lateral (D).

256 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

in the National Museum of Natural History, Smithsonian Institution, Wash-
ington, D.C.; the remaining paratypes in the collection of the author.

Female.—Body (Figs. 1, 2) relatively large. Length 0.54 mm (0.50—0.57
mm) and greatest dorsoventral thickness 0.32 mm (0.30—0.35 mm), based on
10 specimens. Greatest width at level of cephalosome approximately 0.20
mm. Segment of leg 1 fused with cephalosome. Genital segment (Fig. 3)
produced ventrally with concave anterior surface. Greatest diagonal length
of segment 265 um. In ventral view genital segment 143 x 165 um, a little
wider than long (Fig. 4). Abdomen (postgenital segments) small, 80 um long,
and 3-segmented (Fig. 3).

Caudal ramus (Fig. 5) minute, 20 x 20 um, bearing six smooth setae,
longest seta 52 um. Portion of lateral surface of ramus with very small
spinules.

Egg sac (Fig. 6) 240 x 212 um, usually containing three eggs, each egg
approximately 167 x 120 wm. Occasionally only two eggs in egg sac (Fig.
2). Egg sac held between large scooplike fifth legs and anterior concave
surface of genital segment.

Rostrum (Fig. 7) a small prominence. First antenna (Fig. 8) 6-segmented
and 196 um long. Lengths of segments (measured along posterior margin):
31 (39 um along anterior margin), 36, 34, 29, 34, and 24 um respectively.
Armature: 1, 8, 9,3 + | aesthete, 5, and 10 + 1 aesthete. All setae naked.

Second antenna (Fig. 9) 3-segmented and 140 um long not including ter-
minal spines. Exopod 9 x 3.5 wm, bearing two unequal terminal setae. En-
dopod with first segment bearing one seta; second segment carrying two
marginal setae and having terminally four slender setae and two stout slight-
ly clawlike elements with small inner subapical teeth. Inner margin of basi-
pod and first segment of endopod with row of small spinules; similar spinules
along outer margin of second segment of endopod.

Labrum (Fig. 1) prominent. Mandible (Fig. 10) with precoxa armed ter-
minally with a seta and four teeth (two middle teeth with double tips). Coxa-
basis ornamented with crescentic row of spinules, bearing distally two inner
plumose setae and one outer smooth seta. Endopod with three terminal
smooth setae. First maxilla (Fig. 11) with precoxa bearing three spines (two
of them with a subapical tooth) and one small seta. Coxa with one plumose
seta. Basipod with one plumose seta midway along its length and provided
terminally with three plumose setae and a barbed spine. Second maxilla
(Fig. 12) with syncoxa bearing proximally a small rectangular endite with
two setae, followed by a marginal notch and finally by a larger endite bearing
three setae. Basis elongate with a terminal smooth clawlike spine and having
five setae, one very large with unusually long lateral hairs. Maxilliped (Fig.
13) elongate, approximately 255 um including coxa. Endopod, or “‘hand,”’
about 100 wm long. Claw 60 um and bearing one seta and a few spinules
(Fig. 13). Otherwise maxilliped without armature or ornamentation.

VOLUME 94, NUMBER 1 257

VEN

Figs. 8-15. Tegastes acroporanus, female. 8, First antenna, ventral (E); 9, Second antenna,
outer (C); 10, Mandible, anterior (C); 11, First maxilla, posterior (C); 12, Second maxilla,
posterior (C); 13, Maxilliped, inner (D); 14, Claw of maxilliped, outer (C); 15, Leg 1 and
intercoxal plate, anterior (D).

258 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Leg | (Fig. 15) with l-segmented endopod and exopod. Legs 2-4 (Figs.
16, 18, 19) with 3-segmented rami. Formula for armature as follows:

P, coxa 0-0 basis 1-1 exp 2,3
So. lds dk |
P, coxa 0-0 basis 1-0 exp 1-1; 1.3
enp 0-1 il, Z
P, coxa 0-0 basis 1-0 exp I-l 143
enp) 0-3 0-2-5 al,
Pacoxa, 0-0) basise, 1-05 exp lO lel 1e33
enp) s0- ha s0-2< 9 12

Spinelike setae on all four legs with minutely truncate tips. Exopod and
endopod of leg | (Fig. 15) about equal in length. Third endopod segment of
leg 2 in one female with abnormal armature 2, 2 (Fig. 17). Exopod of leg 4
with distal of two inner setae on third segment enlarged, shaped like a
curved blade, with subapical outer teeth (Fig. 19).

Leg 5 (Fig. 20) with greatly enlarged scooplike baseoendopod (Fig. 21)
and elongate slender exopod. Baseoendopod approximately 218 x 130 um,
with an outer smooth seta, a plumose inner seta, and a smooth seta at distal
outer corner. Exopod 105 x 18 um, bearing five naked setae.

Living specimens in transmitted light opaque gray, eye red, eggs gray.

Male.—Body (Fig. 22) resembling in general form that of female. Length
0.47 mm (0.46—0.48 mm) and greatest dorsoventral thickness 0.28 mm (0.28-
0.29 mm), based on 10 specimens. Greatest width at level of cephalosome
approximately 0.18 mm. Genital segment produced ventrally, with sper-
matophore-reservoir divided into anterior foliose lappet concave posteriorly
and posterior rounded lobe separated by smaller lobes (Fig. 23). Abdomen
and caudal ramus similar to female.

Rostrum like that of female. First antenna (Figs. 24, 25) 7-segmented, 161
wm long, and showing strong sexual dimorphism. Lengths of segments
(measured along posterior margin): 33 (36 wm along anterior margin), 31,
17.5, 45, 8, 40.5, and 14 wm respectively. Formula for armature: 1, 8, 6 +
1 aesthete, 9 + | aesthete, 1,3, and 10 + I aesthete. All setae naked. Sixth
segment with swollen proximal part and recurved, almost clawlike distal
part.

Second antenna, labrum, mandible, first maxilla, second maxilla, maxil-
liped, and legs 1-4 similar to those in female. Abnormalities in armature of
legs noted as follows: one male with 1, 2, 1, 1 on endopod of leg 1, and
another male with 2, 3 on third segment of endopod of leg 3.

Leg 5 (Fig. 26) with a single seta on baseoendopod. Free segment elon-
gate, 57 <x 7 wm, bearing two smooth outer marginal setae and two long
unilaterally barbed terminal setae.

VOLUME 94, NUMBER 1 259)

Figs. 16-21. Tegastes acroporanus, female. 16, Leg 2 and intercoxal plate, anterior (D);
17, Third segment of endopod of leg 2, anterior (D); 18, Leg 3 and intercoxal plate, anterior
(D); 19, Leg 4 and intercoxal plate, anterior (D); 20, Leg 5, antero-outer (D); 21, Leg 5, anterior
(B).

260 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Color as in female.

Etymology.—The specific name acroporanus is a combination of the ge-
neric name of the host coral and the suffix -anus, signifying belonging to.

Remarks.—The genus Tegastes Norman, 1903, contains at present 25
valid species. In only 11 of these are both sexes known. In 12 species males
are unknown and in two species females have not been described. Most
species of Tegastes are free-living. However, three species (including the
new species described above) are now known to be associated with inver-
tebrates. Tegastes knoepffleri Médioni and Soyer, 1967, occurs with the
bryozoan Schizobrachiella sanguinea (Norman) at Banyuls in southern
France (Médioni and Soyer, 1967). Tegastes georgei Marcus and Masry,
1970, lives on the scleractinian corals Stylophora sp. and Pocillopora sp.
in the Gulf of Elat, Red Sea (Marcus and Masry, 1970).

The expanded nature of the baseoendopod of the fifth leg in the female
serves to distinguish Tegastes acroporanus from 15 species in which fe-
males are known and where the baseoendopod is not expanded and in some
cases is triangular and tapered distally. These species are: 7. areolatus
Monard, 1935, 7. brasiliensis Jakobi, 1953, T. calcaratus Sars, 1910, T.
clausi Sars, 1904, T. dalmatinus Petkovski, 1955, T. elenae Marcus, 1963,
T. falcatus (Norman, 1868), T. flavidus Sars, 1904, T. grandimanus Sars,
1904, T. longimanus (Claus, 1863), T. nanus Sars, 1904, T. neapolitanus
(Claus, 1863), T. pulcher Pesta, 1932, T. riedli Pesta, 1959, and 7. tenuis
Pesta, 1932. The remaining 10 species may be separated from the new
species on other grounds. In 7. andrewi (T. Scott, 1894) the inner margin
of the hand of the maxilliped has a curved spiniform process and a dilated
appendage. In 7. chalmersi Thompson and A. Scott, 1903, the length of the
female is 0.3 mm and the hand of the maxilliped has spinules along the inner
edge. In 7. georgei Marcus and Masry, 1970, the third segment of the
exopod of leg 4 bears a forked seta. In 7. knoepffleri Médioni and Soyer,
1967, the maxilliped has several spines and a ciliated cup on the second
segment. In 7. minutus Sewell, 1940, the length of the female is 0.29 mm
and the hand of the maxilliped is produced proximally. In 7. perforatus
Lang, 1935, the male first antenna is 8-segmented with the last 3 segments
relatively unmodified. In 7. porosus Petkovski, 1955, the female genital
segment is drawn out into 2 spikes. In 7. satyrus (Claus, 1860) the bas-
eoendopod of the fifth leg in the female is narrow, of about equal length
throughout, and rounded terminally. In 7. sewrati Monard, 1936, the bas-
eoendopod of leg 5 in the female is quadrangular and the hand of the max-
illiped is swollen proximally. In T. edmondsoni Pesta, 1932, the hand of the
maxilliped bears spinules along the inner side.

A close relationship between Tegastes acroporanus and T. georgei 1s
suggested by the nature of leg 5 in the female. 7. acroporanus differs from
the Red Sea species in having 6 apical setae on the endopod of the second

VOLUME 94, NUMBER I 261

Figs. 22-26. Tegastes acroporanus, male. 22, Lateral, with legs 1-4 amputated (A); 23,
Urosome, lateral (D); 24, First antenna, ventral (E); 25, First antenna, dorsal (E); 26, Leg 5,
lateral (E).

262 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

antenna, in having 6 setae (none of them forked) on the third exopod seg-
ment of leg 4, and in showing strong sexual dimorphism in the sixth segment
of the first antenna of the male.

Literature Cited

Claus, C. 1860. Beitraege zur Kenntniss der Entomostraken.—Erstes Heft, pp. 1-28. Mar-
burg.

———. 1863. Die frei lebenden Copepoden mit besonderer Beriicksichtigung der Fauna
Deutschlands, der Nordsee und des Mittelmeeres.—pp. 1-230. Leipzig.

Humes, A. G. 1979. Coral-inhabiting copepods from the Moluccas, with a synopsis of cyclo-
poids associated with scleractinian corals.—Cah. Biol. Mar. 20:77—107.

Jakobi, H. 1953. Neue Tegastiden (Harpacticoida-Copepoda) von der Kueste Santa Catarinas
(Brasilien).—Dusenia 4:173-—180.

Lang, K. 1965. Copepoda Harpacticoidea from the Californian Pacific coast—Kungl. Svenska
Vetensk. Akad. Handl. 10(2):1—560.

Marcus, A. 1963. Tegastes elenae n. sp. harpacticoide nouveau de la Mer Noire.—Vie et

Milieu 14:561—569.

, and D. Masry. 1970. Tegastes georgei n. sp. a new harpacticoid (Crustacea, Cope-

poda), found on corals in the Gulf of Elat.—Israel J. Zool. 19:169-174.

Medioni, A., and J. Soyer. 1967. Copépodes harpacticoides de Banyuls-sur-Mer. 6. Nouvelles
formes associeés a des bryozoaires.—Vie et Milieu, ser. A, 18:317-343.

Monard, A. 1935. Les harpacticoides marins de la région de Salammbo.—Bull. Stat. Océan-

ogr. Salammbo, no. 34, pp. 1-94.

. 1936. Note préliminaire sur la faune des harpacticoides marins d’ Alger.—Bull. Trav.

Stat. d’Aquiculture et de Péche de Castiglione, Alger, pp. 47-85.

Norman, A. M. 1869. Shetland final dredging report.—Part II. On the Crustacea, Tunicata,

Polyzoa, Echinodermata, Actinozoa, Hydrozoa, and Porifera.—Rept. 38th Mtg. Brit.

Assoc. Adv. Sci., 1868, pp. 247-336.

. 1903. New generic names for some Entomostraca and Cirripedia.—Ann. Mag. Nat.

Hist., ser. 7, 11:367—369.

Pesta, O. 1932. Marine Harpacticiden aus dem Hawaiischen Inselgebiet.—Zool. Jahrb. (Syst.)

63:145-162.

. 1959. Harpacticiden (Crust. Copepoda) aus submarinen Hohlen und den benachbarten

Litoralbezirken am Kap von Sorrent (Neapel).—Pubbl. Staz. Zool. Napoli 30 suppl.:95—

Wis

Petkovski, T. K. 1955. Weitere Beitrage zur Kenntnis der Grundwasser-Copepoden der ad-
riatischen Kuste.—Acta Mus. Maced. Sci. Nat. 3:209-225.

Sars, G. O. 1903. An account of the Crustacea of Norway with short descriptions and figures

of all the species. V. Copepoda Harpacticoida. Parts I and II Misophriidae, Longipe-

diidae, Cerviniidae, Ectinosomidae (part), pp. 1-28. Bergen Museum, Bergen.

. 1904. An account of the Crustacea of Norway with short descriptions and figures of

all the species. V. Copepoda Harpacticoida. Parts V and VI Harpacticidae (concluded),

Peltidiidae, Tegastidae, Porcellidiidae, Idyidae (part), pp. 57-80. Bergen Museum, Ber-

gen.

. 1910. An account of the Crustacea of Norway with short descriptions and figures of

all the species. V. Copepoda Harpacticoida. Parts XXIX and XXX Tachidiidae (con-

cluded), Metidae, Balaenophilidae, Supplement (part), pp. 337-368. Bergen Museum,

Bergen.

Scott, T. 1894. Report on the Entomostraca from the Gulf of Guinea, collected by John
Rattray, B.Sc.—Trans. Linn. Soc. London, 2nd ser. zool., 6(1):1—161.

VOLUME 94, NUMBER 1 263

Sewell, R. B. S. 1949. The littoral and semiparasitic Cyclopoida, the Monstrilloida and
Notodelphyoida.—John Murray Exped., 1933-1934, Sci. Repts., 9:17-199.

Thompson, I. C., and A. Scott. 1903. Report on the Copepoda collected by Professor Herd-
man, at Ceylon, in 1902.—Rept. Govt. Ceylon Pearl Oyster Fish. Gulf of Manaar, Suppl.
Repts. 7:227-307.

Boston University Marine Program, Marine Biological Laboratory,
Woods Hole, Massachusetts 02543.

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 264-275

THREE NEW SPECIES OF THE EVIOTA EPIPHANES
GROUP HAVING VERTICAL TRUNK BARS
(PISCES: GOBIIDAE)

Susan J. Karnella and Ernest A. Lachner

Abstract.—Three new species of gobiid fishes of the genus Eviota Jenkins
from Indo-Pacific marine waters are described, namely, E. disrupta, E.
fasciola, and E. irrasa. These species are related to the E. epiphanes group,
sharing a number of meristic and morphological characters, including the
cephalic sensory pore system. They are distinguished from each other and
from E. epiphanes by the presence, size, shape, and intensity of particular
color marks, especially the dark mark or marks on the fleshy base of the
pectoral fin, and the dark vertical bars on the trunk. The salient differences
in the color patterns of preserved specimens of these four allopatric species
are compared and illustrated, and their distributions are plotted on a map.

In this study we treat three new species that resemble Eviota epiphanes
Jenkins in certain color characters and share many meristic and morpho-
logical characters (Group I, Lachner and Karnella, 1980). The shared char-
acters are: complete sensory pore system for Eviota; branching of certain
pectoral rays; male genital papilla nonfimbriate; vertebrae total 26; dorsal-
anal fin ray formula 9/8; pelvic fin rays always I,4; interradial pelvic fin
membrane reduced; segments between consecutive branches of the fourth
pelvic fin ray usually 1; no elongation of the first dorsal fin; and five ventral
midline spots. These four species may superficially resemble each other in
preserved color pattern, depending upon the state of preservation or degree
of development of particular color marks. These similarities may be ob-
served in the following characters: the presence of bars dorsally on the head
and nape; the presence of at least short bars or saddles along the dorsal
midline of the trunk; the head, laterally and ventrally, having large scattered
chromatophores, often arranged in clusters forming spots; caudal peduncle
with a well developed dark subcutaneous spot that is integrated with a weak
to strong subcutaneous vertical bar; and the first dorsal fin with a dark,
irregularly mottled or barred pattern.

We find that meristic and morphometric characters are not useful in dis-
tinguishing these species. The discriminating characters segregating the
species are specific color marks, primarily the type of mark on the fleshy
base of the pectoral fin and the development, uniformity, and intensity of
the vertical trunk bars. Color differences of the 3 new species and E. epi-
phanes are shown in Table 1.

265

VOLUME 94, NUMBER |

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podojaaop
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266 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Methods

The methods of obtaining counts and measurements and the presentation
of these data are given by Lachner and Karnella (1978, 1980). Detailed
analysis of particular color marks was accomplished by recording pertinent
data for each specimen. The specific color marks investigated were the
following: dark mark on fleshy base of pectoral fin, its shape, intensity, if
double or single, and relative size; transverse bars on head, dorsally, and
nape, their intensity, density (whether less dense centrally than marginally),
continuity over dorsal midline, if broken into spots laterally, or more in-
tensely pigmented laterally; dark vertical bars on trunk, their discreteness
(if sharply separated from pale interspaces), uniformity (if complete bars or
broken laterally into vertical, oblong marks), continuity over dorsal midline,
length of bars from dorsal midline ventrally, width of bars relative to pale
interspaces, density of bars, and presence of two narrow bars at base of
caudal fin; presence of pigmentation on the scale pockets, and the integra-
tion of this pigmentation with the trunk bars; presence of scattered chro-
matophores, clusters of chromatophores forming spots, or intermediate con-
ditions, on cheek, opercle and ventral portion of head; dark mark on lower
opercle and branchiostegal membranes, the degree of development and in-
tensity relative to cheek pigmentation; and the nature of the pigmentation
and its intensity on the anal and first and second dorsal fins.

The description of the cephalic sensory pore systems and the cutaneous
papillae systems follow Lachner and Karnella (1978, 1980).

Eviota disrupta, new species
Fig. 1

Material examined.—229 specimens from several localities in eastern
Oceania; total size range 8.1—-16.1 mm; no gravid females found.

Holotype.—USNM 220912, (15.4), male; Society Is., Bora-Bora, south
edge of Passe Teavanui, 6 Apr. 1970, C. L. Smith and R. Mathews, S70-13.

Paratypes.—All paratypes from same collection as holotype, originally
consisting of 181 specimens (AMNH), distributed as follows: USNM
220567, 60 (8.8-16.1). ANSP 143060, 5 (12.9-15.3). AMS 1.21421-001, 5
(11.1-14.8). BPBM 22891, 5 (11.8-15.3). CAS 45395, 5 (11.0-15.4). AMNH
39133, 100 (9.1-16.1).

Nontype material.—SOCIETY ISLANDS: Raiatea: USNM 220571, 1
(11.5). Huahine Nui: USNM 220570, 1 (14.7); AMNH 39134, 1 (14.6), S70-
2; AMNH 39136, 1 (13.1), S70-3. Moorea: CAS 45397, 1 (12.5), sta. 67,
GVF Reg. 1143; USNM 220569, 1 (12.4); CAS 45401, 4 (11.7-15.4), sta. 20,
GVF Reg. 1352; AMNH 39135, 3 (8.1-13.7), S70-64. Maiao: CAS 45409, 2
(12.2, 13.4), sta. 21, GVF Reg. 1353. Tahiti: CAS 45411, 2 (13.3, 15.2), sta.
18, GVF Reg. 1350; CAS 45410, 1 (14.5), sta. 84, GVF Reg. 1160; BPBM

VOLUME 94, NUMBER 1 267

Fig. 1. Eviota disrupta: upper, USNM 220912, holotype, male, 15.4 mm SL, Bora-Bora;
lower, USNM 220567, paratype, male, 15.3 mm SL, Bora-Bora.

8630, 4 (11.5—14.5). Raroia: CAS 45414, 4 (11.5-12.6), sta. 58, GVF Reg.
112. MANGAREVA: BPBM 13587, 2 (13.4, 13.5); BPBM 13556, 1 (14.7);
BPBM 13546, 3 (10.0-11.8); BPBM 13548, 4 (13.9-14.0). SAMOA IS-
LANDS: Tutuila: USNM 220996, 10 (11.2-14.4); BPBM 11309, 1 (12.3).
TONGA ISLANDS: USNM 220565, | (13.2), Te Vega Cr. 7, sta. 301.

Diagnosis.—Base of pectoral fin with two sharply defined, circular dark
spots, clearly separated, of about equal size; pectoral spots intensely pig-
mented, more so than trunk bars; head (dorsally) and nape with large dark
spots, usually arranged transversely and discontinuous along dorsal midline;
trunk with dark, discretely outlined, vertical bars, some of which are almost
always irregularly broken into elongate, oval or oblong shaped marks on the
anterolateral portion of the body; trunk bars extending ventrally below mid-
side of body and often discontinuous over dorsal midline on anterior part
of trunk; scale pockets on trunk not pigmented.

Description.—Dorsal fin VI-I,8(1), VI-I,9(/4); anal fin I,7(1), I,8(/4); pec-
toral fin 16(4), 17(/1); pelvic fin I,4(/4); fourth ray of pelvic fin with 8-11
branches, average 9.4; number of segments between consecutive branches
of fourth pelvic fin ray 0-2, average 1.1; pelvic fin membrane reduced;
branched caudal fin rays 11(1), 12(3), 13(4), 14(1); segmented caudal fin rays
16(1), 17(/4); scales highly deciduous, lateral scale rows 23(1), 24(1), 25(1);
transverse scale rows 8(1); breast scaleless.

268 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Spinous dorsal fin not elongate. Depressed pelvic fin rarely extending to
origin of anal fin, never beyond.

Cephalic sensory pore system is pattern 1; cutaneous papillae system is
pattern A.

Male genital papilla not fimbriate.

Vertebrae 10(7) precaudal, 16(7) caudal, total 26(7).

Color in preservative.—This species is similar in general color pattern to
E. fasciola and E. irrasa except for the following characters: the pectoral
base almost always has two discrete dark circular spots, the upper one
larger or equal to the lower spot, and the spots are the most intense body
pigmentation; in a few specimens the pectoral spots are weakly connected
by less dense chromatophores; first and third transverse nape bars anterior
to the dorsal fin origin are frequently separated along the midline forming
spots laterally, the second bar often short and continuous over dorsal mid-
line; trunk bars discretely outlined, irregularly interrupted anteriorly form-
ing elongate vertical marks, and more regular posteriorly; bars usually ex-
tending below lateral midline, and sometimes discontinuous over dorsal
midline; trunk bars sometimes with less dense or pale interiors; dark spot
on lower opercle and branchiostegal membrane is conspicuous and often
divided into two parts or nearly so; a small, sixth subcutaneous ventral
midline spot may be present at the base of the procurrent caudal fin rays,
but is never associated with a subcutaneous bar.

Color in life. —The following observations on color of a living specimen
from Samoa (USNM 220996) were recorded by R. Wass: body greenish-
yellow, operculum bluish-green, bars and spots brown.

Geographic distribution.—This species occurs in eastern Oceania (Figure
6) from the Samoa and Tonga Islands eastward to Mangareva.

Etymology.—The specific epithet, disrupta is Latin, meaning ““broken

9)

up,’ in reference to the pattern of the vertical trunk bars.

Eviota fasciola, new species
Figs. 2 and 3

Material examined.—345 specimens from the Great Barrier Reef, Aus-
tralia, and numerous localities in western Oceania; total size range 7.8—19.1
mm; smallest gravid female 13.6 mm SL.

Holotype. —USNM 220560, (18.0), male; One Tree Island, Queensland,
Australia, 22 Nov. 1966, V. G. Springer, 66-7.

Paratypes.—All paratypes were collected from the Great Barrier Reef,
Queensland, Australia. ONE TREE ISLAND: USNM 220564, 25 (10.2-
19.1); CAS 45396, 5 (12.5—17.7); WAM P26843-001, 5 (11.2—16.8); previous
three lots from same collection as holotype. USNM 220563, 31 (11.4—18.0),
VGS 66-16; AMS I.15641-049, 1 (17.5). HERON ISLAND: LACM 32820-8,

VOLUME 94, NUMBER 1 269

Fig. 2. Eviota fasciola: upper, USNM 220560, holotype, male, 18.0 mm SL, One Tree
Island; lower, USNM 220559, paratype, female, 13.5 mm SL, Lizard Island.

2 (16.4-16.5). ENDEAVOUR REEFF (collected by J. Tyler and C. L. Smith
in 1969): ANSP 143059, 4 (13.5-15.3), TS,A-17; ANSP 143058, 11 (10.5-
15.4), TS,A-13; ANSP 143057, 1 (12.4), TS,A-6; AMNH 39132, 1 (13.6),
$69-19; LIZARD ISLAND (collected by Australian Museum team in 1975):
AMS 1.21422-001, 15 (10.2—17.6), DFH 75-14; AMS [I.19473-177, 103 (9.3-
16.9), LZ 75-60; USNM 220559, 11 (11.8-16.5), LZ 75-70; USNM 220955,
3 (10.4-12.4), LZ 75-17; AMS I.19483-071, 76 (10.2-17.8), LZ 75-71.

Fig. 3. Eviota fasciola, ANSP 143059, paratype, male, 15.3 mm SL, Endeavour Reef.
(Drawn by J. R. Schroeder)

270 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Other material.—PALAU ISLANDS: CAS 45407, 1 (12.1), Ngadarak
Reef, sta. 106, GVF Reg. 612. ULITHI ATOLL: CAS 45400, 7 (11.5—13.8),
sta. 178, GVF Reg. 975; CAS 45403, 10 (10.1-13.4), sta. 181, GVF Reg.
978; CAS 45412, 4 (11.5-12.9), sta. 179, GVF Reg. 976. IFALUK ATOLL:
CAS 45402, 2 (11.9-12.5), sta. 76, GVF Reg. 379; CAS 45406, 1 (11.3), sta.
7, GVF Reg. 128; CAS 45413, 1 (9.1), sta. 109, GVF Reg. 230. MARIANAS
ISLANDS: Saipan: CAS 45405, 7 (11.1-14.9), sta. 47, GVF Reg. 833; CAS
45399, 14 (9.2-16.0), sta. 46, GVF Reg. 832. USNM 149916, 13 (8.9-14.6);
USNM 150022, 1 (12.0). Guam: CAS 45398, 8 (8.6-14.7), sta. 59-28, GVF
Reg. 1856; UG 4512, 10 (8.7-15.6). AMS 18403-001, 2 (14.1, 15.3). MAR-
SHALL ISLANDS: Enewetak Atoll: AMS [I.18405-001, 1 (14.7); AMS
18401-003, 1 (14.5); USNM 220558, 2 (12.9, 13.7); BPBM 8241, 2 (13.5, 14.8);
USNM 220562, 3 (13.3-14.7); LACM W63-281-2, 1 (14.8); LACM 6679-61,
6 (8.7—15.3); USNM 220568, 4 (10.4—15.5); LACM W63-283-4, 2 (10.9, 14.5).
Majuro Atoll: AMS 1.18399-001, 1 (14.0). GILBERT ISLANDS: Abaiang
Atoll: AMS 1.18052-001, 15 (8.9-15.5). TROBRIAND ISLANDS: AMS
I.17102-052, 1 (15.4); USNM 220561, 2 (10.6, 14.2).

Tentative identifications. —-KAPINGAMARANGI ATOLL: CAS 45408,
10 (7.8-11.9), sta. 62, GVF Reg. 365; CAS 45404, 2 (9.1, 9.5), sta. 8, GVF
REG, SI

Diagnosis.—Base of pectoral fin with large dark kidney shaped to semi-
circular mark; intensity of pectoral mark greater than that of trunk bars;
head (dorsally) and nape with dark transverse bars, usually complete over
dorsal midline, occasionally one or more bars broken up into spots laterally,
and about one-half of the specimens with spots or bars darker laterally than
dorsally; trunk with prominent dark, discrete vertical bars, descending to
ventral portion of body and continuous over dorsal midline; scale pockets
on trunk not pigmented.

Description.—Dorsal fin VI-I,8(3), VI-I,9(/5); anal fin I,8(/7), I,9(1); pec-
toral fin 16(7), 17(11); pelvic fin I,4(/9); fourth ray of pelvic fin with 6-12
branches, average 9.1; number of segments between consecutive branches
of the fourth pelvic fin ray 0-2, average 1.1; pelvic fin membrane reduced;
branched caudal fin rays 11(3), 12(6), 13(3); segmented caudal fin rays
17(17); lateral scale rows 23(1), 24(5S), 25(/); transverse scale rows 6(3);
breast scaleless.

Spinous dorsal fin not elongate. Depressed pelvic fin usually not extending
to origin of anal fin, never beyond it.

Cephalic sensory pore system is pattern 1; cutaneous papillae system is
pattern A.

Male genital papilla not fimbriate.

Vertebrae 10(9) precaudal, 16(8), 17(1) caudal, total 26(8), 27(1).

Color in preservative.—Specimens from Australia almost always have an
intensely pigmented kidney-shaped spot, rarely a semicircular spot, extend-

VOLUME 94, NUMBER | 7} I

ing over the height of the pectoral base, the upper portion usually wider.
The pectoral spot is the most prominent mark on the body. In a few of the
type specimens the pectoral spot is more heavily pigmented on the upper
and lower portions than the middle portion, and rarely the mark is divided
into upper and lower spots on specimens from Lizard Island. A pale area
almost always lacking chromatophores is present on anterior part of base
of pectoral fin and posterior portion of opercle. A dark circular spot occurs
on the upper portion of the mesial surface of base of the pectoral fin. Head
and trunk with prominent vertical dark bars, usually wider than the pale
interspaces; 12—13 trunk bars from origin of spinous dorsal fin to midpe-
duncular spot, extending from dorsal midline to ventral portion of trunk,
the last bar overlying the dark subcutaneous peduncle spot. Usually
a small dark spot bordering upper procurrent rays just posterior to last bar
and two narrow vertical bars at base caudal fin, the posterior one on bases
of caudal fin rays; spot and bars are often undeveloped. The trunk bars may
be reduced in size and intensity in some specimens, or they may have pale
interiors, and the trunk pigmentation occasionally may be obscure, but the
head and pectoral base marks persist. Three dark transverse bars on nape
anterior to the origin of the spinous dorsal fin; a fourth, the anteriormost
bar on the occipital area, usually present but often more irregular than other
bars on nape and sometimes segmented into patches laterally and dorsally
along the midline. A dark triangular mark is present immediately above and
behind the eyes. One or more of the nape bars may be broken into spots
and/or intensely pigmented laterally. Cheek with 6—7 large patches of chro-
matophores, a large diffuse patch on anterior part of opercle and a heavily
pigmented patch on lower margin of opercle and overlapping branchiostegal
membrane. Cheek, opercle and branchiostegal patches may be irregularly
arranged in some specimens. Lower part of head and chin with patches of
chromatophores similar to those on cheek. Five subcutaneous spots on ven-
tral midline from origin of anal fin to end of caudal peduncle, associated
with subcutaneous bars on lower part of trunk; bars absent from upper part
of trunk. The fifth spot and lower subcutaneous bar are continuous with the
caudal peduncle spot. Fins of the Australian specimens often pale, lacking
prominent coloration. Spinous dorsal fin pale or with a diffuse dusky oblique
band through the lower portion, the anterior basal portion, as well as the
outer half, pale or dusky to brown, the margin sometimes darker. Second
dorsal and anal fins dusky to pale. The remaining fins may have the rays
bordered with fine chromatophores, otherwise they are pale.

The specimens from Trobriand Islands have a semicircular spot at the
base of the pectoral fin, the anterior margin of which is nearly vertical. The
nape and trunk bars may be discontinous along the dorsal midline.

Specimens from western Oceania, from the Palau to the Gilbert Islands
(Fig. 6), have a more variable shaped pectoral spot, ranging from semicir-

DL, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

cular or oblong to somewhat kidney shaped; a specimen from the Gilbert
Islands has the upper and lower portions of the spot enlarged with a central
constriction, suggesting a two-spotted condition. The pectoral spot is less
intense than in the Australian specimens, often equal in intensity to the
trunk bars.

The only sexual dimorphism in coloration observed was that the females
may be slightly paler.

Geographic distribution.—This species occurs in western Oceania from
the Palau to the Gilbert Islands, the Trobriand Islands, and the Great Barrier
Reef (Fig. 6).

Etymology.—tThe specific epithet is taken from the Latin word fascia, in
reference to the bands nearly encircling the trunk.

Eviota irrasa, new species
Figs. 4 and 5

Material examined.—7 specimens from the Philippine Islands; total size
range 12.3-17.6 mm; no gravid females found.

Holotype. —USNM 220566, (16.0), male; Philippine Is., Palawan Prov.,
Cocoro I. (Cuyo Is.); 25 May 1978, V. G. Springer and Smithsonian team,
SP 78-26.

Paratypes.—USNM 220954, 5 (12.3-17.6); data as for holotype. USNM
220582, 1 (17.4), Philippine Is., Palawan Prov., Putic I. (Cuyo Is.), 22 May
1978, V. G. Springer and Smithsonian team SP 78-18.

Diagnosis.—Base of pectoral fin with two dark circular to oval spots, not
sharply defined and often weakly connected, the upper spot somewhat larger
than the lower; pectoral spots moderately pigmented, the intensity about
equal to trunk bars; head (dorsally) and nape with dark, transverse bars
continuous over dorsal midline, and somewhat more heavily pigmented lat-
erally; trunk with regular dark, vertical bars, somewhat diffuse in outline,
continuous over dorsal midline and usually not extending below midside
anteriorly and much shorter posteriorly on peduncle; scale pockets dorso-
laterally on trunk often weakly pigmented.

Description.—Dorsal fin VI-I,9(7); anal fin I,8(7); pectoral fin 16(3), 17(4);
pelvic fin I,4(7); fourth ray of pelvic fin with 8-11 branches, average 9.9;
number of segments between consecutive branches of fourth pelvic fin ray
1-2, average 1.2; pelvic fin membrane reduced; branched caudal fin rays
12(1), 13(1); segmented caudal fin rays 16(1), 17(6); lateral scale rows 24(2);
transverse scale rows 6(1); breast scaleless.

Spinous dorsal fin not elongate. Depressed pelvic fin rarely extending to
origin of anal fin, never beyond.

Cephalic sensory pore system is pattern 1; cutaneous papillae system is
pattern A.

VOLUME 94, NUMBER 1 213

Fig. 4. Eviota irrasa, USNM 220566, holotype, male, 16.0 mm SL, Cuyo Islands. (Drawn
by J. R. Schroeder)

Male genital papilla not fimbriate. Vertebrae 10(6) precaudal, 16(6) caudal,
total 26(6).

Color in preservative.—This species is similar in the general color pattern
to E. fasciola and E. disrupta. It differs in the following characters: E.
irrasa has upper and lower dark, circular to oval, pectoral base spots that
may be joined by scattered chromatophores, the upper spot somewhat larg-
er; nape bars rarely broken up into spots; trunk bars mostly confined to
dorsolateral portion, only the anterior bars extending to ventral area, the
bars becoming shorter posteriorly and occurring only dorsolaterally; trunk
bars sometimes diffuse, the pigmentation becoming irregular and outlining
some of the scale pockets; dorsal and anal fins darker than in E. fasciola,
the oblique band in the spinous dorsal fin dusky to black, wide, and some
Specimens with small intense black areas on membrane in middle portion
of fin just posterior to first spine and in an oblique row following the next
3-4 spines; trunk bars may extend into basal portions of first and second
dorsal fins; anal fin dusky to blackish.

Fig. 5. Eviota irrasa, USNM 220954, paratype, female, 14.0 mm SL, Cuyo Islands.

274 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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E. epiphanes represented by shaded area AANMUAANNIUUUII

Fig. 6. Geographic distributions of Eviota fasciola, E. disrupta, E. irrasa, and E. epi-
phanes, based on material examined.

Geographic distribution.—Known only from the Cuyo Islands, Palawan
Province, Philippine Islands (Fig. 6).

Etymology.—The specific name irrasa is Latin, meaning unpolished or
unshaved, and is used in reference to the indiscrete nature of the pigmen-
tation pattern of the head and trunk compared with E. fasciola and E.
disrupta.

Discussion

Although Eviota epiphanes, E. disrupta, E. fasciola, and E. irrasa are
superficially similar in many aspects, certain color marks are consistently
different. The salient differences among these species are summarized in
Table 1. No evidence of intergradation of these species is shown in our
data. The four species occur in distinct geographical areas and are not so
far known to be sympatric (Fig. 6). Because of the poor state of preservation
of the specimens from Marcus Island (BPBM 7154, 23 spec.; BPBM 7155,
4 spec.) we identify these only tentatively as E. epiphanes.

Acknowledgments

We thank the curatorial and support staff at the following institutions for
the loan and exchange of specimens pertinent to this study: Academy of

VOLUME 94, NUMBER 1 275

Natural Sciences of Philadelphia (ANSP), American Museum of Natural
History, New York (AMNH), Australian Museum, Sydney (AMS), Bernice
P. Bishop Museum, Honolulu (BPBM), California Academy of Sciences,
San Francisco (CAS), Los Angeles County Museum of Natural History
(LACM), and the University of Guam (UG). Our thanks also go to Richard
C. Wass, Office of Marine Resources, Government of American Samoa,
who donated a collection of Eviota disrupta from Samoa. We are especially
indebted to C. L. Smith (AMNH) for allowing the generous exchange of
type specimens of Eviota disrupta to various institutions, including the de-
position of the holotype at the National Museum of Natural History, Wash-
ington, D.C. (USNM). The drawings (Figs. 3 and 4) were rendered by J. R.
Schroeder.

The manuscript was reviewed by C. E. Dawson who provided valuable
suggestions.

Literature Cited

Lachner, E. A., and S. J. Karnella. 1978. Fishes of the genus Eviota of the Red Sea with
descriptions of three new species (Teleostei: Gobiidae).—Smithsonian Cont. Zool.
286122:

, and . 1980. Fishes of the Indo-Pacific genus Eviota with descriptions of eight
new species (Teleostei:Gobiidae).—Smithsonian Cont. Zool. 315:1—127.

Department of Vertebrate Zoology (Fishes), National Museum of Natural
History, Smithsonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 276-294

TAXONOMY AND POSTMARSUPIAL DEVELOPMENT
OF A DOMINANT DEEP-SEA EURYCOPID
ISOPOD (CRUSTACEA)

George D. Wilson

Abstract.—Eurycope iphthima n. sp. is a valuable subject for studies of
systematics and population biology in abyssal Crustacea because of it nu-
merical dominance, size, and robustness. In this introductory paper, the
species is described, and developmental variation is discussed for the fol-
lowing characters: pereopod VII, antennula, rostrum, uropods, oostegites,
and male pleopods I and II. Some evidence indicates this species is facul-
tatively hermaphroditic and its sexual development is partly independent of
somatic growth.

Because deep-sea sampling is so difficult, the biology of few abyssal ben-
thic Crustacea is known well enough to examine problems of ontogenetic
and population variation. Yet a knowledge of variability is critical to a clear
understanding of the systematics of a group. In order to make these types
of analyses in an important deep-sea group, the Eurycopidae, this paper
introduces Eurycope iphthima new species, a dominant abyssal isopod. This
species has a large body allowing its retention by standard sampling tech-
niques: the smallest post-marsupial individuals are greater than one milli-
meter in length. Of additional value, E. iphthima has a very robust body,
insuring undistorted specimens that are relatively easy to measure. Fur-
thermore, E. iphthima has potential ecological importance to deep-sea fau-
nal assemblages because of its numerical dominance in some areas. In two
large epibenthic sled samples collected during Woods Hole Oceanographic
Institution (WHOIJ) cruise no. 106 (stations 326 and 328), this species is the
most abundant isopod, accounting for 23% of the total individuals. This
introductory paper describes Eurycope iphthima n. sp. and the post-mar-
supial development of a number of its systematically important characters.
Subsequent papers will discuss the taxonomy, population biology, and zoo-
geography of this important species.

Terminology.—Hessler (1970) reviewed the terminology used for the de-
velopment of asellote isopods and concluded that the most informative sys-
tem is that of Zimmer (1926), introduced for cumacean development. In this
paper, the updated system of Hessler (1970) is used, except that the juvenile
Stage is not directly associated with instars due to this species’ complicated
development (discussed below). Because three preparatory female stages

VOLUME 94, NUMBER 1 277

before the brooding stage are recognized (Fig. 9D—-F), the number after
preparatory refers to the developmental stage, rather than the number of
times the animal has entered that condition (as in Hessler, 1970), e.g., pre-
paratory 3 female means the third and final preparatory female intermolt
stage.

For structures on pleopods I and II of the male, the reader is referred to
the excellent paper by Veuille (1978a), whose descriptive terms are used
here with the exception that instead of his term, copulatory whip, stylet is
used (Fig. 10K) as in past papers (Hessler, 1970). The small, thin distal
projection on the protopod of the male pleopod II (Fig. 10K) is called the
vermiform appendage after Menzies (1962).

Measurements.—Measurements are made as described in Wilson and
Hessler (1980), with the following additions. Overhang is the ratio be-
tween the length of the rostral projection from the frons (Fig. 8D—a), and
the rostral length (Fig. 8D—b). The length of the medial lobe of the anten-
nular article one (Fig. 7C—ml) is measured from the insertion of article two
to the lobe’s distal tip. In the male pleopod I length—width ratio, the width
is measured at the proximal edge of the dorsal orifice (Fig. 10K—do).

Family Eurycopidae
Subfamily Eurycopinae
Eurycope iphthima, new species
Figs. 1-5

Holotype.—Preparatory 3 female, 7.0 mm long, deposited in the United
States National Museum (USNM), catalog number 181106.

Paratypes.—10 individuals, USNM 181107; 5 individuals, American Mu-
seum of Natural History, New York, catalog number 16253; 5 individuals,
Zoological Museum of the University of Copenhagen; 273 individuals re-
tained in the collection of Robert R. Hessler, Scripps Institution of Ocean-
ography.

Type-locality.—WHOI station 326, R/V Chain cruise no. 106, 22 August
1973, 50°04.9-05.3’N, 14°23.8-24.8’W, 3859 meters, estimated bottom tem-
perature (from Fuglister, 1960) 2.5°C.

Derivation of name.—phthima is Greek, meaning strong or stalwart, re-
ferring to the robust nature of the species.

Other material.—See Table 1.

General distribution.—Abyssal Central North Atlantic and the rise and
slopes of Northeastern Atlantic, 2379-5000 meters.

Diagnosis.—Body length approximately 2.1 times width, cephalon widest
at level of pereopod I coxae. Rostrum (Fig. 8) almost always with some
overhang, especially when rostrum narrow, width varies around 0.1 ceph-
alon width, with u- or v-shaped anterior notch. Female and immature an-

278 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

aos
\\
SAN

oy,
Ce

OLSS

Fig. 1. Eurycope iphthima: A-E, I: holotype preparatory 3 female. A, Body, dorsal view;
B, Body, lateral view; C, pereonites 1—4 and associated coxae, cuticular ridges shown only on
pereonite 3; D, rostrum, left oblique view; E, Cephalon, dorsal view; F, Preparatory 2 female,
cephalon, oblique frontal view, antennula and antenna removed on left side; G-H, Precopu-
latory adult male dorsal and lateral views; I, Cephalon, lateral view.

tennular third article (Fig. 7B) approximately same length as second article;
basal article medial lobe slightly shorter than article 2; copulatory male
antennular third article (Fig. 7B) slightly shorter than second article, basal
article medial lobe around same length as second article, flagellar articles

VOLUME 94, NUMBER I 24)

Table 1.—Localities for Eurycope iphthima n. sp.

Mean Mean Number
west north Depth in of Number of
Station latitude longitude meters samples individuals
*WHOI 326
(type locality) $0°0S’ 14°24’ 3859 1 294
*WHOI 80 34°49' 66°34’ 4970 1 3
WHOI 83? 34°46’ 66°30’ 5000 1 1 juv.
WHOI 121? 35°49’ 65°10’ 4800 1 2 frags
WHOI 287? IS BGs 54°52! 4957 1 9 frags
WHOI 288 11°03’ 55°04’ 4423 1 it
WHOI 318 DOM TE 13°20’ 2506 1 105
WHOI 321 50°12! ISe36y 2879 1 150
WHOI 323 50°08’ IB 35 27 3347 1 86
WHOI 328 SO205% 15°45’ 4431 1 261
WHOI 330 50°43’ SZ’ 4632 1 35
WHOI 334 40°43’ 46°14’ 4400 1 200
* Allen 50 43°46’ 3°38’ 2399 1 19
tINCAL 1.8 55°01’ ZO" 2890 4 18
INCAL 2.1 SOM: 132" 2641 4 108
INCAL 2.2 48°19’ e231 4829 1 12
INCAL 2.3 47°30’ 9°34’ 4284 11 96
INCAL 2.4 46°00’ 10°21’ 4802 1 1

* More information on these stations in Wilson and Hessler (in press).
+ Expedition Intercalibration, described in Sibuet (1979).

after article 8 wider than long. Pleopod I of copulatory male (Fig. 10I—K)
length 3.6 width, dorsal orifice position 0.68 medial length; distal tip truncate
in ventral view, tapered medially, lateral lobes thin to ventrally-curving
spines closely adpressed to medial lobe; medial lobe thick, rounded medi-
ally, thinning laterally to dorsal shelf. Pleopod II of copulatory male (Fig.
10K) with long vermiform process, 0.15 total pleopod length. Uropodal en-
dopod length less than protopod width, exopod length greater than half
endopod length (Fig. 9A—C).

Additional description and variation.—Body robust, natasome broadly
rounded in dorsal view, vaulted, muscular. Lengths of mature adults greater
than 4 to 5 mm, largest individuals may exceed 7 mm; females larger than
males, median lengths (ranges) 5.5 mm (4.1-—7.4) vs. 4.9 mm (4.0—6.3); mean
stage 4 length 2.2 mm, standard deviation 0.1, range 2.0-2.5.

Rostral variability common (Fig. 8): median rostral length-cephalic width
ratio 0.19, range 0.18—0.22; medial rostral width-length ratio 0.36, range
0.22—0.64; median overhang 0.28, range 0.0—0.33; rostrum rarely juts abrupt-
ly from frons; when rostrum narrow, overhang more distinct; median values
vary from population to population.

280 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

“B.C Die

Fig. 2. Pereopods: A, Pereopods I-IV, III-IV intact, dactylus IV enlarged; B, Pereopod
I with oostegite, dactylus enlarged; C—E, Left natapods from precopulatory male: pereopod
V, whip seta from ischium enlarged; pereopod VI; pereopod VII. Plumose setal rows indicated

by basal insertions and single representative setae.

VOLUME 94, NUMBER I 281

Fig. 3. Left mandibles: A-E, Preparatory 2 female. A, Dorsomedial view; B, Palp, lateral
plan view; C, Medial view, plan view of triturative surface; D, Spine row, ventral view; E,
Lacinia mobilis and incisor, posterior view, f—posterior fold in ventral tooth; F—H, Copulatory
male. F, Dorsal view; G, Lateral view, palp in plan view; H, Setation of palp distal tip; I-J,
stage 4 individual. I, Lateral view; J, Dorsai view.

Pereonite 7 medial length much greater than pereonite 5 or 6, pereonite
7 bulla enlarged and rounded; pereonites 5 and 7 subequal.

Antennula of copulatory male (Fig. 7D) with about 60 articles, length
beyond proximal 2 articles 4.0 times length of article 1 medial margin; female
antennula (Fig. 7B) values, 26 articles and 2.7 length ratio; basal article
medial lobe with 5—7 unequally bifid setae in adults.

Left mandibular incisor (Fig. 3) with 3 large teeth, ventral tooth with fold
in posterior surface (Fig. 3E—f), dorsal tooth with small tooth on dorsal
side. Lacinia mobilis with 6 teeth on medial surface grading into 2—3 den-
ticles on dorsal surface. Spine row with 7-9 spines having teeth or spinules

i)

N

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

J

ag, yp

XN

Appendages from preparatory females: A, Maxillula; B, maxilla with enlargement

Fig. 4.
of mediobasal denticles and setae; C—E, Maxilliped. C, Ventral view, epipod detached, setae

not shown; D, Distal tip of endite, fan setae bases shown with examples to either side; E,
Setation on palp articles 3 and 4; F, Pleopod II, ventral view, hemiplumose setae shown on
left side, cuticular ridges shown only on apex and distal tip; G, Pleopod II, lateral view; H,

Pleopod III; I, Pleopod IV; J, Pleopod V.

VOLUME 94, NUMBER 1 283

and very fine accessory setae on anteriormost 4-6 spines. Molar approxi-
mately same length as mandibular condyle; triturative surface oval, teeth on
posterior and dorsal margins low and rounded, 6-8 setae on posterior mar-
gin. Mandibular palp shorter than mandibular body; male palp longer than
female palp: male palp article 2 length 0.44 times mandible body length,
female 0.38.

Maxilliped (Fig. 4C—E) in adults with 6—9 receptaculi, around 7 short thick
unequally bifid setae on ventral face of coxa and basis; palp merus width
1.2 times endite width. Epipod broad with subquadrate lateral expansion,
with 2-4 short thick setae on ventromedial ridge, length 1.8 times width.

Pereopod-body length ratios in pereopods I, III, IV: 0.74, 0.90, 1.05.
Bases of pereopods I-II of copulatory male longer than in preparatory 3
female: copulatory male basis-body length ratios in first 4 pereopods 0.21,
0.20, 0.20, 0.22; for preparatory female 0.18, 0.19, 0.20, 0.20. Male pereo-
pods (Fig. 2C—E) V—VII propodus width 0.51 times length, ventral setal row
only on distal half of margin. Dactylus of pereopod VI largest of natatory
limbs, pereopod VII dactylus smallest. Only one large seta at proximal
posterior angle of pereopod VII basis.

Pleopod II of both sexes (Figs. 10K and 4F, G) with strongly recurved
lateral margins. Male pleopods I (Fig. 10I—-K) widest at insertion, with paired
row of 9-10 hemiplumose setae. Male pleopod II (Fig. 10K) length 1.7 times
width, depth 0.3 times length, with about 14 hemiplumose setae on each
lateral margin. Stylet 0.69 times pleopod length. Female pleopod II (Fig.
4F-G) length 0.79 times width, depth 0.3 times length, with about 11 hem-
iplumose setae on lateral margins.

Uropodal protopod (Fig. 9A—C) strongly broadened medially, width great-
er than length, ventral and medial margins with dense row of long, thin
unequally bifid setae.

Remarks.—Eurycope iphthima belongs to a speciose subgroup within the
genus wherein the differences between the member species are small but
definitive. Other described species in this subgroup are E. complanata Bon-
nier 1896, E. hanseni Ohlin 1901, E. inermis Hansen 1916, and E. ratmanovi
Gurjanova 1946. These species are being revised and new species are being
described in research now in progress.

Eurycope complanata is most similar to E. iphthima and occurs within
the same geographic area although at much shallower depths. E. compla-
nata 1S narrower in the body (I/w ratio 1.9) and has a narrower, less over-
hanging rostrum. Additionally, the antennular article 3 is distinctly longer
than article 2 and the endopod of the uropod is longer than the width of the
protopod.

Potential synonyms.—Menzies (1962, pp. 141-142) describes 4 individuals
of Eurycope complanata collected by the R/V Vema in Lamont Geological
Observatory Biotrawl no. 7 (5779 m 29°17.6’N, 57°20.3’W). The large male

284 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

( ,;

fi "GS

LP

< 4 yy a = Ps
E 6) Z s

i? a

IR

Fig. 5. Development of pereopod VII: Pereopod VII shown darkened in right lateral oblique
view; basis of pereopod VI included for comparison; only representative setae in plumose setal
rows shown. A, Manca 1; B, Manca 2, arrow indicates pereopod anlagen below cuticle; C,
Manca 3; D, Stage 4; E, Stage 6.

figured by Menzies appears to be E. iphthima, but this cannot be determined
with certainty because this material seems to be missing from the collections
of the American Museum of Natural History (H. S. Feinberg, pers. comm.).
Chardy (1979) lists E. complanata as occurring at stations 2000 meters and
deeper in the Bay of Biscay. The present data and other information show
that E. iphthima occurs in the Bay of Biscay in this depth range and E.
complanata occurs shallower, 860—1920 meters (Bonnier, 1896; Wolff, 1962;
unpublished data).

Postmarsupial Development

The postmarsupial development of deep-sea asellote isopods has been
reviewed by Wolff (1962) and Hessler (1970). More recently Thistle and
Hessler (1977), and Haugsness and Hessler (1979) provide developmental
data for 2 abundant eurycopids. This section will describe the development
of several taxonomically important characters and discuss the relationship
between somatic growth and sexual maturation in Eurycope iphthima and
in the species studied by the above authors.

Pereopod VII.—The first 5 stages can be recognized on the basis of the

VOLUME 94, NUMBER |

Nw
oO
Nn

2 29

- “=
2 T

:

16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 50 52 54 56 58
0.2mm Size Classes

Fig.6. Representative body length-frequency histogram of all intact individuals from WHOI

318. Stages three to five, 3—5. Juveniles, J. Precopulatory and copulatory, PC&C. Preparatory

one, P/. Preparatory three, P3. Brooding, B. Individuals outside of the 1.7—5.7 mm range and
preparatory two females appeared in other samples.

development of the seventh pereopod and pereonite (Fig. 5). Manca 1 has
no apparent pereopod VII and the corresponding segment appears dorsally
as a triangular plate having no lateral margins. The manca 2 pereopod is a
subcuticular primordium and the pereonite is somewhat larger but still with
no lateral margins. In manca 3 the pereopod is external but rudimentary

286 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 7. Development of the antennula: A, Stage 4 individual; B, Preparatory 2 female; C,
Precopulatory male, enlargement showing setal detail, distance ml illustrates medial lobe
length; D, Copulatory male, not all aesthetascs present.

with no functional joints or setae. At this stage the seventh pereonite be-
comes part of the lateral margin but the anterolateral corner is not pointed.
Similar to Jaera a. albifrons Bocquet (1953) the stage 4 pereopod VII be-
comes functional but is distinctly smaller, with fewer setae compared to the
anterior natapods. In most specimens where the distal parts of the limb have
been lost in sampling, stage 4 can be recognized by a basis which is smaller
and much less robust than the basis of pereopod VI. In stage 5S the basis is
the same length as pereopod VI but slightly less robust. In stage 6 and
larger, pereopods VII and VI bases are of approximately equal size and
robustness.

Instar size. —Figure 6 shows a representative size-frequency histogram
from WHOIT 318 which illustrates most of the instars discussed below. Eu-
rycope iphthima increases in length by roughly | mm in 2 molts. This in-
crease is less for mancas (with these stages falling between 1 and 2 mm) and
perhaps greater for older individuals. Mancas 1—3 were approximately 1.1,
1.4 and 1.8 mm respectively. Stage 4 averaged 2.2 mm, varying only one or
two tenths in either direction. Significantly, the mean length for stage 4,
which was well represented at the localities studied in detail, did not vary

VOLUME 94, NUMBER 1 287

Fig. 8. Development and variation of the rostrum: Top illustrations, left oblique dorsal
view; bottom illustrations, dorsal view; cuticular ridges shown. A, Stage 4 individual, 2.2 mm;
B, Male, 5.1 mm; C, Preparatory 3 female, 6.7 mm; D, Preparatory | female, 5.8 mm, distance
a is the rostral projection from the frons, distance b is the rostral length; E, Male, 6.5 mm; F,
maie, 5.1 mm. A—D, are common conditions; E-F are rare.

geographically. Stage 5 mean length at one abundant station (WHOI 318,
Fig. 6) is 2.7 mm (SD = 0.14, N = 27, range = 2.4—2.9). Beyond stage 5
the molts are difficult to recognize, although at some localities they appear
as modes in the size-frequency distribution. Sexual development cannot be
used to recognize specific stages because it appears to be independent of
size. Stage 6 is about 3.2—3.6 mm and stage 7 is at approximately 4.0-4.4
mm. Sexual maturity may occur at any stage after 7, the earliest being stage
8, roughly 4.4-5.0 mm. Beyond this stage the data do not allow a clear
association between instar and size.

Antennula.—Besides the sexual pleopods, dimorphism is most apparent
in the development of the antennula (Fig. 7). In mancas and stage 4, the
flagellum has only a few articles, no aesthetascs and a more rounded medial
lobe having 4 or 5 unequally bifid setae. One taxonomically diagnostic char-
acter, article 3 being subequal or shorter than article 2, is stable, even in
these stages. As development proceeds, more articles are added to the fla-
gellum with copulatory males achieving 50-60 articles and preparatory 3
females with only 20-25. Notably the females have aesthetascs only on the
distal articles while in males they may occur on the full length of the fla-
gellum. Male flagellar articles are wider than long and female articles are
longer than wide. However, in spite of the shorter articles, the male anten-
nula is longer. With increasing instar number the medial lobe becomes slight-
ly more pointed and the distal setae increase to 6 or 7.

288 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

6000 i ———_ =

(TE
Sate a

Fig. 9. A—C, Uropoda: A, Stage 4 individual; B, Preparatory 2 female; C, Copulatory male,
posterior view. D-F, Developing oostegites: D, Preparatory | female; E, Preparatory 2 female;
F, Preparatory 3 female.

Rostrum.—In spite of the variability associated with the rostrum, there
are some developmental components that can be recognized (Fig. 8). In
mancas and stage 4, the rostrum is fairly broad: rostral-cephalic width ratio
around 0.15. With maturity, the rostrum narrows to a ratio of around 0.10,
although in some males this can be as small as 0.05. The rostral-frons over-
hang can be seen to increase with age, ranging from 0.0 to 0.12 in the earliest
stages to 0.10-0.33 at maturity. Mature males also tend to have greater
overhang values than females. Between-individual variability in rostral
shape also seems to increase with age such that all the manca and stage 4
individuals within a sample appear very similar while adults can be sub-
stantially different.

Uropoda.—On the uropod (Fig. 9A—C), key taxonomic characters remain
relatively unchanged through the life cycle: the endopod is always shorter
than the width of the protopod, and the exopod length is always greater
than half the endopod length. However, in the mancas the rami are thicker
than in the adults: the endopod-protopod width ratio is 0.35; in mature adults
this decreases to around 0.31 and 0.24 for males and females respectively.
Note that the males retain some width; they also have endopods that are

VOLUME 94, NUMBER I 289

slightly longer than females. The protopodal row of long, thin unequally
bifid setae increases allometrically with size: in the manca 3, it has around
8—9 setae; in adults there are 20—22 setae, with no apparent dimorphism in
this or other setal characters.

Oostegites and female maturation.—Four separate stages of development
of the oostegites could be recognized in Eurycope iphthima (Fig. 9D-F).
Wolff (1962) previously recognized a great variation in the sizes of devel-
oping oostegites in several eurycopids and munnopsids, but could not relate
them to a developmental sequence for lack of data. The material of E.
iphthima clearly shows that the increase in the size of the rudimentary
oostegites is correlated with somatic growth and is a direct consequence of
approaching maturation in females. The female juvenile phase, that is, in-
dividuals with no evidence of oostegite buds but not mancas, always spans
stages 4 and 5, although it clearly goes into stages 6, 7 or larger in some
individuals. Therefore one can find outwardly ‘juvenile’ females from 2.0 to
an observed maximum of 5.6 mm. It is not known whether the large ‘ju-
veniles’ represent totally non-reproductive individuals or females that have
brooded once. These largest females are at least one stage larger than the
smallest brooding females.

Preparatory | females are characterized by rudimentary oostegites which
appear as small buds on the medial edge of the coxae of the first 4 pereopods
(Fig. 9D). The tissue below these buds appears to be somewhat more dense
and granular, thus aiding their identification. There was some interindividual
Variation in size of the buds but none approach the size of the next stage.
Preparatory | females occurred at lengths from 3.5 mm (stage 6) up to 5.8
mm, well beyond stage 9.

A second rare state occurs where the rudimentary oostegite is approxi-
mately the same size as the coxa (Fig. 9E). Only one or two preparatory 2
females were found at each station, varying in length from 4.9 to 7.0 mm.
The rarity of this state may be due to most females passing directly from
preparatory 1 to preparatory 3 in one molt.

The preparatory 3 oostegite is much larger than the coxa but not extending
to the midline (Fig. 9F). This condition was found in females as small as 4.6
mm (stage 8 or 9) to as large as 7.5 mm. The females in this state should be
considered as being fully copulatory and sexually mature, because this is the
final stage before brooding. In support of this assertion, Veuille (1978b) has
shown that in Jaera the final preparatory females receive the males. Fur-
thermore, several preparatory 3 females of E. iphthima which were dis-
sected proved to have apparent cuticular organs and gonads full of oocyte ma-
terial. In this state the oostegites were observed to vary from being thin,
transparent lamellae to fat opaque pads. That these latter oostegites are in
a premolt condition is shown by the presence of developing structures under
the old cuticle.

290 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 10. Development of male pleopods I and II. A—C, Stage 4 individuals, all 2.2 mm long;
D, Stage 5, 2.6 mm; left pleopod II detached to show developing stylet; E, Stage 7, 3.7 mm;
F, Juvenile stage, 4.6 mm; G—H, Precopulatory male 4.4 mm; G, Pleopods separated; H,
Enlargement of pleopod I distal tip, spines on lateral lobes straight, setae omitted from left
side; I-K, Copulatory male, 5.4 mm, setae if omitted shown by their insertions; I-J, Pleopod
I distal tip, lateral (Showing curved spine) and ventral views, m/—medial lobe, //—lateral lobe;
K, Ventral view, pleopods separated, enlargement shows hemiplumose seta typically found in
rows on venter of pleopod I and lateral margin of pleopod II, st—stylet, va—vermiform ap-
pendage, do—proximal edge of dorsal orifice.

In the final brooding condition, the oostegites become quite large and
form the typical isopodan brood chamber. The oostegal lamella is very thin
with an axial thickening for stiffness, and with several groups of fine sensory
setae on the external surface. Brooding females were fairly rare in the col-
lections and ranged in size from 4.5 to 5.8 mm. Because preparatory 3
females longer than 7 mm were found, it is reasonable to suppose that the

VOLUME 94, NUMBER 1 291

brooding female can attain this size. The size-spread in mature and brooding
females indicates that this species may reproduce at least twice in a life
cycle. If this is the case for the eurycopid species examined by Wolff (1962),
his data on oostegite and body size are easily explainable within the frame-
work of the present study.

Male pleopods I and IT.—Bocquet (1953) demonstrated that the male
pleopod development in Jaera is not rigidly associated with particular in-
stars. Similarly, these sexual characters of male Eurycope iphthima
show considerable variation in their development. Two types of variation
were recognized: the stage at which the pleopod begins to develop, and the
degree of development of the pleopods at any one instar. Pleopod I appears
in the manca 3, as in other deep-sea asellotes (Wolff, 1962; Hessler, 1970;
Haugsness and Hessler, 1979), or later in stage 4 or 5. Males with developing
pleopods were found over a broad range of sizes (2.0—4.9 mm). If the total
Variation in the development of pleopod I is considered, a gradual matura-
tion is seen, although for each individual the pleopod growth is no doubt
saltational. Consequently, it was not possible to classify the stages of male
maturation other than to note when a male was fully copulatory.

At first appearance the first pleopod is a flat, bifurcate lobe of variable
size above an undifferentiated pleopod II (Fig. 10A—C). In the degree of
development shown in Fig. 10C the primordia of the endopod and exopod
of pleopod II can be seen underneath the cuticle. As the instars proceed,
the first pleopod extends to its full length relative to the pleotelson, and the
second pleopod separates with the endopod and exopod becoming external,
but undeveloped (Fig. 10D). Hemiplumose setae (Fig. 10K) appear in rows
on the lateral margins of pleopod II and the ventral surface of pleopod I.
The penile papillae enlarge and enter the anterior funnel of the first pleopods
while the medial lobe separates from the lateral lobe and becomes more
setose. At this point the stylet is fully elongated and its sperm duct is com-
plete and open. During this development the distal tip of pleopod II becomes
produced into a thin projection of variable length, the vermiform process,
which in copulatory males (Fig. 10K) is nearly as long as the exopod. The
pleopod I lateral lobe changes in the final molt from a straight spine (Fig.
10H) to a ventrally-curving spine (Fig. 10I).

Anomalous sexual development.—At WHOI 328 one 4.9 mm long indi-
vidual was distinctly larger than stage 7, but had a developing male pleopod
I on top of an undifferentiated female-like pleopod II. The second largest
male (6.0 mm) at WHOI 326 was observed to have an antennula intermediate
in development between mature males and females. Several of the largest
males from these collections are only in the precopulatory stage, based on
pleopod morphology, while smaller males are fully copulatory. Similar ob-
servations are found in other deep sea species. Hessler (1970) reports a very
large preparatory male of Chelator vulgaris which exhibits feminine-like

292 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

features, and the large illustrated male of Eurycope ratmanovi Gurjanova
(1946) has immature pleopods. These anomalies may well represent mani-
festations of protogynous hermaphroditism, wherein a female may go
through one brooding cycle and then changes into a male. Both in E. iphth-
ima and in C. vulgaris, the unusual males are the same size or larger than
the smaller modes of brooding females. Protogynous hermaphroditism has
been reported for other janiroideans (Wolff, 1962) and is indicated by adult
intersex individuals of Haploniscidae, Ishnomesidae, Pleurogoniidae, and
other Eurycopidae (unpublished data). These latter individuals had both
developing male pleopods and female characters, either oostegites or cuticu-
lar marks of the egg-laying organ (see Veuille, 1978b). It is important to
note that some past reports of intersexes in deep-sea isopods, i.e., Haplonis-
cus percavix Menzies (1962), are probably juvenile developing males. In
spite of this possible confusion, the hermaphroditic condition may be much
more prevalent in deep-sea isopods than previously thought. This certainly
has proved to be the case in tanaids of the family Neotanaidae (Gardiner,
1975).

Discussion.—Hessler (1970) notes the development of most desmoso-
matid species appears to be rigidly scheduled as in Haploniscus biscuspis
Wolff (1962). The size-frequency data on Bellibos buzwilsoni Haugsness and
Hessler (1979) also seems to agree with this. However, Thistle and Hessler
(1977), reporting on Betamorpha characta Hessler and Thistle (1975), de-
scribe some variability in the form of the immature male pleopods I and II,
and could not discern any development of these male pleopods in the manca
3 stage.

Somatic growth in E. iphthima, at least for the earliest stages, is rigidly
determined; stage 4 individuals always fall within a few tenths of a millimeter
of 2.2 mm, regardless of the locality. In contrast to this, the stages of sexual
development for both males and females have broad size ranges over several
localities, clearly spanning a number of instars. Because the observed range
of instars for levels of reproductive development is more restricted within
localities, one possible function of this flexibility may be to allow a deme
of the widespread E. iphthima more readily to adapt to local environmental
conditions.

The results on this species indicate that there are 2 components to de-
velopment that need to be considered: somatic growth, described as the
succession of instars; and sexual development, described as the appearance
of primary and secondary sexual characters. In most desmosomatids, the
appearance of sexual features is closely coupled with somatic growth, while
in E. iphthima and in the genus Jaera there is some independence between
these two components.

However, observed variation of sexual development may require consid-
eration of two different phenomena, decoupled sexual development, and

VOLUME 94, NUMBER 1 293

hermaphroditism. Thus, Chelator vulgaris is an example of rigidly coupled
sexual and somatic development with protogyny, Jaera shows some de-
coupling of sexual growth, and Eurycope iphthima displays characteristics
of both influences on development.

Acknowledgments

This research was conducted in the laboratory of Dr. Robert R. Hessler,
Scripps Institution of Oceanography, from whose collection the material of
Eurycope iphthima was drawn. This latter gentleman and Dr. David Thistle,
Florida State University, reviewed the manuscript and offered a number of
valuable suggestions for its improvement. Mr. Bryan Burnett skillfully inked
the pencil illustrations. Drs. H. S. Feinberg, American Museum of Natural
History in New York, and Torben Wolff, Zoological Museum of the Uni-
versity of Copenhagen, aided in the locating of material of a related species,
E. complanata. The Woods Hole Oceanographic Institution deep-sea ben-
thic ecology program, directed by Dr. Howard Sanders and Dr. Fred Gras-
sle, and the Centre National de Tri d’Oceanographie Biologique, directed
by Dr. Michel Segonzac, are the original sources of the specimens studied.
A National Science Foundation grant to Dr. Robert R. Hessler, DEB77-
24614, provided the funding for this research. I sincerely thank the above
gentlemen and institutions for their involvement in bringing this work to
fruition.

Literature Cited

Bocquet, C. 1953. Recherches sur le polymorphisme naturel des Jaera marina (Fabr.) (Iso-
podes Asellotes).—Arch. Zool. Exp. Gen. 90:187—450.

Bonnier, J. 1896. Edriophthalmes.—Ann. Univ. Lyon 1895:527-689.

Chardy, P. 1979. Structure of deep-sea asellota assemblages in the Bay of Biscay; relationships
with the abyssal environment.—Ambio Special Report, no. 6, pp. 79-82.

Fuglister, F. 1960. Atlantic Ocean atlas of temperature and salinity profiles and data from the
International Geophysical Year of 1957—1958.—Woods Hole Oceanographic Inst. Atlas
Ser. 1:209 pp.

Gardiner, L. 1975. The systematics, postmarsupial development, and ecology of the deep-sea
family Neotanaidae (Crustacea: Tanaidacea).—Smithsonian Cont. Zool. no. 170, 265 pp.

Gurjanova, E. 1946. New species of Isopoda and Amphipoda from the Arctic Ocean.—Comp.
Results, Drifting Exp., Icebreaker “‘Sedov,’’ 1937-1940. Moscow 3:272-297.

Hansen, H. 1916. Crustacea Malacostraca IHI.—Danish Ingolf Exped. 3(5):1—262.

Haugsness, J., and R. Hessler. 1979. A revision of the subfamily Syneurycopinae (Isopoda:
Asellota: Eurycopidae) with a new genus and species (Bellibos buzwilsoni).—Trans. San
Diego Soc. Nat. Hist. 19(10):121-151.

Hessler, R. 1970. The Desmosomatidae (Isopoda, Asellota) of the Gay Head—Bermuda Tran-

sect.—Bull. Scripps Inst. Oceanography 15:1—185.

, and D. Thistle. 1975. On the place of origin of the deep-sea isopods.—Marine Biology

327155165:

Menzies, R. 1962. The isopods of abyssal depths in the Atlantic Ocean.—Abyssal Crustacea.
Vema Res. Ser. 1:79-206.

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Ohlin, A. 1901. Arctic Crustacea collected during the Swedish Arctic Expeditions 1898 and
1899 under the direction of Prof. A. G. Wathorst. I. Leptostraca, Isopoda, Cumacea.—
Bih. Svensk. Vetensk. Aka. Handl. 26, 4(12):1—54.

Sibuet, M. 1979. Biologie. Connaissances générales sur les communautes benthiques abyssales
dans |’ Atlantique nord-est—CNEXO 3:1-96.

Thistle, D., and R. Hessler. 1977. A revision of Betamorpha (Isopoda; Asellota) in the world
ocean with three new species.—Zool. J. Linn. Soc. 60:275-295.

Veuille, M. 1978a. Biologie de la reproduction chez Jaera (Isopode Asellote) I.—Structure

et fonctionnement des pieces copulatrices males.—Cah. Biol. Marine 19:299-308.

. 1978b. Biologie de la reproduction chez Jaera (Isopode Asellote) II.—Evolution des

organes reproducteurs femelles.—Cah. Biol. Marine 19:385-395.

Wilson, G., and R. Hessler. 1980. Taxonomic characters in the morphology of the genus
Eurycope (Crustacea, Isopoda) with a redescription of E. cornuta Sars, 1864.—Cah.
Biol. Marine 21:241-263.

Wolff, T. 1962. The systematics and biology of bathyal and abyssal Isopoda Asellota.—Gal-
athea Rep. 6:1—320.

Zimmer, C. 1926. Northern and Arctic invertebrates in the collection of the Swedish State
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Scripps Institution of Oceanography A-002, La Jolla, California 92093.

PROC. BIOL. SOC. WASH.
94(1), 1981, pp. 295-313

THE AMPHIPOD GENERA EFOBROLGUS AND EYAKIA
(CRUSTACEA: PHOXOCEPHALIDAE) IN
THE PACIFIC OCEAN

J. Laurens Barnard and Charline M. Barnard

Abstract.—Eobrolgus now contains 3 species, spinosus (Holmes) (type),
chumasi n. sp. and pontarpioides (Gurjanova); E. spinosus appears to be
a west Atlantic species introduced into lagoons and estuaries of the north-
eastern Pacific, whereas E. chumasi is a native eastern Pacific species and
the possible ancestor of spinosus. Now that spinosus has returned to the
Pacific, gene flow is perceived to occur with its parent chumashi.

Eyakia has 4 species, robusta (Holmes) (type) (? = ochotica (Gurjano-
va)), calcarata (Gurjanova), uncigera (Gurjanova) and subuncigera (Ku-
drjaschov). The first 2 are redescribed.

Introduction.—The genera Eobrolgus and Eyakia were recently carved
out of the polytypic Paraphoxus Sars by J. L. Barnard (1979). The species
of these genera are now presented and revised.

Of particular interest is the discovery of a new cryptic species of Phox-
ocephalidae near Eobrolgus spinosus (Holmes) (formerly Paraphoxus spi-
nosus) which appears to be an endemic east Pacific species and which points
to the possibility that east Pacific populations of E. spinosus are introduc-
tions from the Atlantic Ocean.

The taxonomy of American and North Pacific Phoxocephalidae is much
more subtle than that found for Australian taxa by Barnard and Drummond
(1978), so that generic distinctions are much finer and fewer.

The generic information of Barnard (1979) is not repeated here so as to
save space for redescription of those species for which good material is
available.

We thank Elizabeth B. Harrison, Janice M. Clark, Irene F. Jewett and
Roland H. Brown for their help with this project. Dr. Janet Haig of the
Allan Hancock Foundation kindly loaned us materials for this study. We
thank John Chapman and Dr. James T. Carlton for their advice about in-
troduced species.

Eobrolgus J. L. Barnard, 1979

This genus differs from Australian brolgins and Paraphoxus by the fol-
lowing factors: (1) proximal spines of article 4 on antenna 2 are thick and
present (in brolgins they are thin or absent); (2) distal spines in the same

296 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

locality are few; (3) inner plate of maxilla 1 with 4 setae (2 in other mentioned
groups); (4) mandibular molar with 4+ spines (3 or fewer in brolgins); (5)
inner plate of maxilliped with one main apical spine (none in brolgins); (6)
maxillipedal dactyl with apical spine fused (not fused in brolgins except in
Elpeddo); (7) epimeron 3 with 3+ long setae (3 or fewer usually short setae
in brolgins); (8) right lacinia mobilis apical branch narrow or absent.

Key to the Species of Eobrolgus

1. Each lobe of telson with 2 apical spines ............ E.. pontarpioides
=Each lobe of telson) with one apical spimels.. +: .4h44.00 7 fee oeaeee y
2. Epimeron 3 naked ventrally, right lacinia mobilis simple in female,
outer plate of maxilla 1 with 9 spines ................... E.. spinosus
— Epimeron 3 with 1-2 ventral setae, right lacinia mobilis bifid in fe-
male, outer plate of maxilla 1 with 11 spines ........... E. chumashi

Eobrolgus spinosus (Holmes)
Fig. If

Paraphoxus spinosus Holmes, 1905:477—478, fig. [unnumbered]; Kunkel,
1918:76-78, fig. 13; Shoemaker, 1925:26-27; J. L. Barnard, 1959:18;
1960:243-249, pls. 29-31; 1961:178; 1964a:105; 1966a:89; 1969a: 197-198;
1969b:224; Reish and Barnard, 1967:18; Bousfield, 1973:125, pl. 34.1.

Description of female ‘‘f’ of Atlantic Ocean.—Head about 18 percent of
total body length, greatest width about 75 percent of length, rostrum un-
constricted, broad, short, reaching middle of article 2 on antenna |. Eyes
large, mostly occluded with pigment, ommatidia ordinary. Article 1 on pe-
duncle of antenna 1 about 1.35 times as long as wide, about 2.1 times as
wide as article 2, ventral margin with about 9 setules, strongly produced
dorsal apex with 2 setules, article 2 about 0.6 times as long as article 1, with
2 apicoventral rows of 4 and 2 setae and 3 lateral setae, primary flagellum
with 7 articles, about 0.55 times as long as peduncle, bearing one short
aesthetasc each on articles 3-6, accessory flagellum with S articles. Antenna
2 thin, not ensiform but article 1 enlarged, article 3 with one anterodorsal
setule plus normal facial formula; spine formula on article 4 = 1-3-4, dorsal
margin with notch bearing one seta and one spine, ventral margin with 5
groups of 2 long to medium setae, one ventrodistal long spine, article 5
about 0.8 times as long as article 4, facial spine formula = 2, dorsal margin
bearing one apical set of short setae, ventral margin with 4 sets of 2 long to
short setae, 3 ventrodistal long to medium spines set facially, flagellum as
long as articles 4—S of peduncle combined, with 8 articles.

Epistome with weak anterior knob. Mandibles with medium palpar hump,
right incisor with 3 normal teeth, left incisor with 3 hump-teeth in 2 branch-

VOLUME 94, NUMBER 1 297

wMl

Fig. 1. Eyakia robusta (Holmes), m = male ‘‘m,”’ w = female ‘‘w,’’ y = female “‘y.”’ Eob-
rolgus chumashi, new species, e = female ‘‘e.”’ Eobrolgus spinosus (Holmes), f = female ‘‘f.”’
Capital letters = A, antenna; F, accessory flagellum; L, lacinia mobilis; M, mandible; N, molar;
P, pereopod; lower case letters to left of capitals refer to specimens described in text; lower
case letters to right of capitals or freely placed in figures are as follows: b, broken; 1, left; r,
right; s, setae removed.

es, right lacinia mobilis simple, pointed, left lacinia mobilis with 4 teeth plus
One accessory tooth, middle teeth not shortened, right rakers 6, left rakers
6 plus one rudimentary, molars composed of short bulbous protrusions,
right molar with 4 primarily medium spines plus one short seta strongly
disjunct, left molar with 4 primarily medium spines plus one short seta
strongly disjunct, each molar with plume, palp article 1 short, article 2 with
2 long and short inner apical setae and one other short inner seta, article 3
about as long as article 2, oblique apex with 7-8 spine setae, basofacial

298 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

formula = 0. Each outer lobe of lower lip with one cone. Inner plate of
maxilla | ordinary, bearing one long apical pluseta, one similar apicomedial
seta, 2 apicolateral much shorter setae, outer plate with 9 spines, palp thin,
scarcely biarticulate, article 2 with one apical spine, one apicolateral and 2
medial setae. Plates of maxilla 2 extending equally, outer not broader than
inner, outer with 2 apicolateral setae, inner with 2 medial setae. Inner plate
of maxilliped with one large thin apical spine, 2 apicofacial setae, 4 medial
setae, outer plate with 8 medial and apical spines, no apicolateral setae,
palp article 1 without apicolateral seta, article 2 without lateral setae, medial
margin weakly setose, article 3 with 4 facial setae, 2 lateral setae, nail of
article 4 fused, with 2 accessory setules.

Coxa | scarcely expanded apically, anterior margin straight, main ventral
setae of coxae 1-4 = 8-8-8-5, posteriormost seta of coxae 1—4 slightly short-
ened, anterior and posterior margins almost parallel, posterior margin con-
vex, posterodorsal corner rounded, posterodorsal margin short, V-shaped,
width-length ratio = 18:19. Gnathopods generally ordinary, width ratios on
articles S—6 of gnathopods 1-2 = 25:36 and 27:39, length ratios = 65:73 and
53:71, palmar humps ordinary, palms strongly oblique, article 5 of gnatho-
pod | elongate, ovate, posterior margin rounded-flat, long, article 5 of
gnathopod 2 ovate, posterior margin rounded, short, almost produced.

Pereopods 3-4 similar to each other, facial setae formulas on article 4 =
4 and 3, almost parallel to apex, on article 5 = 4 and 3, main spine of article
5 extending to M. 100 on article 6, article 5 with no proximoposterior spines,
spine formula of article 6 = 4+ 5 and 4+ 5 plus middistal seta, some
spines especially long, acclivity on inner margin of dactyls of pereopods 3-—
4 absent, emergent setule short, almost fully immersed, midfacial pluseta
ordinary. Coxae 5-7 posteroventral setule formula = 2-1-1. Articles 4—5 of
pereopods 5-6 narrow, facial spine rows sparse, facial ridge formula of
article 2 on pereopods 5—7 = 0-1-1. Width ratios of articles 2, 4, 5, 6 on
pereopod 5 = 53:33:30:13, of pereopod 6 = 68:26:16:9, of pereopod 7 =
80:17:16:8, length ratios of pereopod 5 = 78:32:40:38, of pereopod 6 =
90:56:45:43, of pereopod 7 = 100:23:22:27, article 2 of pereopod 7 reaching
middle of article 4, posterior margin with 8 tiny serrations, only medial apex
of article 6 finely and sparsely combed, bearing 1—2 weak digital processes.

Posteroventral corner of epimeron | rounded, posterior margin straight,
naked, anteroventral margin with 2 medium setae, posteroventral face with
one long seta. Posteroventral corner of epimeron 2 rounded, posterior mar-
gin weakly convex, naked, facial setae = 5, no pair set vertically. Postero-
ventral corner of epimeron 3 rounded, posterior margin straight, serrate,
setose (4), with one dorsal setule notch, ventral margin naked.

Urosomite 1 with lateral setule at base of uropod 1, no ventral setae,
articulation line short, urosomite 3 unprotuberant dorsally. Rami of uropods

VOLUME 94, NUMBER 1 Dy)

1-2 with articulate but tightly fixed apical nails, outer ramus of uropod 1
with 3 dorsal spines, inner with 2, outer ramus of uropod 2 with 2 dorsal
spines, inner with one dorsomedial spine, peduncle of uropod | with 4 ba-
sofacial setae and one apicolateral spine, medially with 4 marginal spines,
apicalmost enlarged but not displaced. Peduncle of uropod 2 with 5 thin
dorsal spines, medially with one medium apical spine, apicolateral corner
of peduncles on uropods 1-2 with comb. Peduncle of uropod 3 with 4 ventral
spines, dorsally with 2 lateral spines, one medial setule, rami feminine, inner
extending to M. 50 on article 1 of outer ramus, apex with one long seta,
medial and lateral margins naked, article 2 of outer ramus elongate, 0.25,
bearing 2 short to medium setae, medial margin of article 1 with 4 setae,
lateral margin with 3 acclivities, spine formula = 2-2-2-2, setal formula =
0. Telson especially long, length-width ratio = 35:29, almost fully cleft, each
apex of medium width, rounded, scarcely protruding, lateral acclivity shal-
low, narrow, bearing ordinary lateral setule, spine next medial longer than
setule; midlateral setules diverse, largest of medium size.

Young male ‘‘p’’ of Pacific Ocean (assumed transplant).—Like Atlantic
spinosus in (1) presence of 9 spines on outer plate of maxilla 1; (2) simple
right lacinia mobilis; (3) coxa 4 setose; (4) uropods 1—2 with comb; (5) rami
of uropod 2 with one spine each; (6) apical setae on outer ramus of uropod
1 medium in length; (7) apical nails on rami of uropods 1-2 large; (8) no
ventral setae on epimeron 3.

Female ‘‘u’’ of Pacific Ocean, possible hybrid.—Like Atlantic spinosus
in items of paragraph above: 2, 3 (but coxa 4 with only | seta), 4, 5 (but
only outer ramus with spine), 6, 7, 8; but outer plate of maxilla | with 11
spines like chumashi.

Male ‘‘b.’’—Article 2 of antenna | with 6-7 narrowly spread ventral setae,
primary flagellum with 9 articles, one calceolus each on articles 1—5. Facial
spine formula on article 4 of antenna 2 = 3-4-1, of article 5 = 2, latter with
4 dorsal sets of male setae and no calceoli, ventrodistal apex with 2 thin
spines, flagellar calceolus formula = 2,3,5,7,9...n.

Basofacial setal formula of article 3 on mandibular palp = 0-1, article 2
with 4 apical setae. Outer plate of maxilla 1 with 9 spines.

Coxa 4 more elongate than in female, width-length ratio = 4:5. Pereopods
3-4 thinner than in female; article 2 of pereopods S—7 narrower than in
female; article 5 of pereopod 7 with 1—2 male setae.

Epimera 1-3 broadened, posterior margin of epimeron 3 not shortened,
setal formulae, epimeron | anteroventral = 2, posteroventral = 1, epimeron
2 facial = 3-4, epimeron 3 posterior = 4, facial = 0, ventral = 0. Spine for-
mulae of uropods, uropod | peduncle apicolateral = 1, basofacial = 4, uro-
pod 2 peduncle dorsal = 5, dorsal spines on outer ramus of uropod 1 = 2,
of uropod 2 = 2, inner ramus of uropod 1 = 1, of uropod 2 = 1, ventral

300 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

spines on peduncle of uropod 3 = 5, spine formula on article 1 of outer
ramus = 1-2-2-2-2, setal formula = 1-1-1-1-1, setae on outer ramus of article
2 as in female.

Telson elongate, length-width ratio = 4:3, distal spines shortened, for-
mula = setule, spine, setule, each lobe with long single denticle row.

Voucher material.—Male **b’’ 4.37 mm and male ‘“‘c’’ 4.02 mm, Woods
Hole, Massachusetts, USA, 9 July 1875, surface, A.E.V. [Verrill]; female
“f°? 4.58 mm, Amityville [Long Island, New York, USA], 6 July 1938, Acc.
no. 149428; young male “‘p’’ 3.22 mm (no antenna 2), Friday Harbor, Wash-
ington, USA, 8896, 21 October 1920; female “‘u’’ 1.88 mm, Newport Bay,
California, USA, station 50, 1964 January (see J. L. Barnard, 1959).

Remarks.—One presumes that this is an Atlantic species that has been
introduced into estuaries, bays and lagoons of the northeastern Pacific
Ocean, such as Friday Harbor and Newport Bay (and Morro Bay, San
Francisco Bay, etc.) where it closely matches the endemic Pacific E. chu-
mashi. Female *‘u’’ cited above is disturbing because it has 11 spines on
the outer plate of maxilla 1 like chumashi yet has the spinosus morphology
in other characters. Study on hybdridization is warranted. The two morphs
may have been separated by Pleistocene closure of the Panamic isthmus
and now have been rejoined by mankind without complete genetic isolation.
Other explanations can be visualized and need not be discussed until the
morphs are explored further.

Barnard (1960) did not detect this problem because finely drawn charac-
ters as now used had not been introduced. Most of the Allan Hancock
Foundation materials will require reidentification, a task we leave to some-
one who wants to pursue the introduction problem.

Despite the amazing superficial resemblance of this species to Paraphox-
us oculatus, the two taxa cannot be congeneric because of (1) the mandib-
ular molar spine counts; (2) the spine distribution on article 4 of antenna 2;
(3) the short article 1 of the mandibular palp; (4) the oblique apex of palp
article 3 on the mandibular palp; (5) the non-foliate right lacinia mobilis; (6)
the presence of 4 setae on the inner plate of maxilla 1; (7) the stout apical
spine on the inner plate of the maxilliped; (8) the presence of basal arma-
ments on article 5 of pereopods 3-4; (9) the presence of long setae on
epimeron 3; and, in addition, the lesser values of setae on article 2 of antenna
1 not being fully apicad and the dactylar nail of the maxilliped being fused.

This species is also confusable with Foxiphalus obtusidens but the fol-
lowing characters distinguish spinosus from obtusidens: (1) the apicad po-
sition of the setae on article 2 of antenna 1; (2) the short head; (3) the non-
ensiform antenna 2; (4) the simple right lacinia mobilis; (5) the thin apex of
the palp on maxilla 1; (6) the tightly fixed apical nail on the inner ramus of
uropod 1; (8) the lack of ventral setae on the urosome.

A few west American samples of *‘Paraphoxus spinosus’’ have been reex-

VOLUME 94, NUMBER 1 301

amined and some found to be the new species, Eobrolgus chumashi, a
cryptic species similar to spinosus. But most of the material in the Allan
Hancock Foundation needs to be reidentified and we have not examined the
material from the Gulf of California, Carmen Island, identified by Shoe-
maker (1925) which is presumably in the American Museum of Natural
History.

The open-sea character of chumashi and the estuarine-lagoon-shallow
open-sea character of spinosus suggest the latter has been introduced onto
the Pacific Coast from the western Atlantic. But more investigation of the
two species is required.

Distribution.—Western Atlantic Ocean from Vineyard Sound to North
Florida (fide Bousfield, 1973), depths unknown, no samples in Smithsonian
collections with stated depths; eastern Pacific Ocean, confirmed localities,
Friday Harbor and Newport Bay (lagoon), shallow water; much material to
be reidentified in Allan Hancock Foundation.

Eobrolgus chumashi, new species
Fig. le

Description of female ‘‘e.’’—Body dwarfed. Head about 21 percent of
total body length, greatest width about 67 percent of length, rostrum un-
constricted, broad, short, almost reaching apex of article 2 on antenna 1.
Eyes large, mostly occluded with pigment, ommatidia especially large. Ar-
ticle 1 of peduncle on antenna | almost 1.5 times as long as wide, about
twice as wide as article 2, ventral margin with 7-8 setules, produced dorsal
apex with 2 setules, article 2 about 0.55 times as long as article 1, with 2
apicoventral rows of 4 and | setae and 2 lateral setae, primary flagellum
with 7 articles, about 0.8 times as long as peduncle, bearing one short aesthe-
tasc each on articles 3-6, accessory flagellum with 5 articles. Antenna 2 not
ensiform, article 3 dorsally naked but with normal facial setules, spine for-
mula of article 4 = 1-3-5, dorsal margin with notch bearing 2 setae and one
spine, ventral margin with 5 groups of 2 long to medium setae, one ven-
trodistal long spine, article 5 about 0.72 times as long as article 4, facial
spine formula = 3, dorsal margin bearing one set of small setae apically,
ventral margin with 4 sets of 1—2 long to short setae, 3 ventrodistal long to
medium spines set facially; flagellum about 0.95 times as long as articles 4—
5 of peduncle combined, with 7 articles.

Epistome unproduced. Mandibles with weak to medium palpar hump,
right incisor with 3 teeth, left incisor with 3 hump-teeth in 2 branches, right
lacinia mobilis bifid, distal branch not shorter than proximal, distal branch
broad, proximal branch narrow, both branches simple, pointed, left lacinia
mobilis with 4 teeth, middle teeth shortened, right rakers 7 plus 2 rudimen-
taries, left rakers 7 plus 3 rudimentaries, molars composed of elongate bul-

302 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

bous plaques, right molar with 5 primarily medium spines plus one short
spine strongly disjunct, left molar with 6 primarily medium spines, no spine
disjunct, each molar with plume, palp article 1 short, article 2 with one short
inner apical seta and 2 other short inner setae, article 3 about 1.1 times as
long as article 2, oblique apex with 9-7 spine-setae, basofacial formula =
0. Each outer lobe of lower lip with one cone. Inner plate of maxilla 1
ordinary, bearing one medium apical pluseta, one similar apicomedial seta,
2 apicolateral shorter setae, outer plate with 11 spines, no cusps, palp article
2 with one apical spine, one apicolateral, 2 apicomedial and 3 submarginal
setae. Plates of maxilla 2 extending equally, outer broader than inner, outer
- with 3 apicolateral setae, inner with 2 medial setae. Inner plate of maxilliped
with one large thick apical spine, 2 apicofacial setae, 3 medial setae, outer
plate with 3 medial spines set apicad tightly, no apicolateral setae or cusp,
palp article 1 without apicolateral seta, article 2 with one apicolateral seta,
medial margin of article 2 moderately setose, article 3 with 4 facial setae,
one lateral seta, nail of article 4 fused, with 2 accessory setules.

Coxa | scarcely expanded apically, anterior margin straight, main ventral
setae of coxae 1-4 = 7-8-7-0, posteriormost seta of coxae 1-3 shortened,
anterior and posterior margins parallel, posterior margin almost straight,
posterodorsal corner sharp-rounded, posterodorsal margin short, concave,
V-shaped, width-length ratio of coxa 4 = 13:17. Gnathopods ordinary,
width ratios on articles 5-6 of gnathopods 1-2 = 25:33 and 26:33, length
ratios = 65:56 and 49:52, palmar humps small, palms oblique, article 5 of
gnathopod | elongate, ovate, posterior margin rounded-flat, long, article 5
of gnathopod 2 ovate, posterior margin rounded, short, almost produced.

Pereopods 3-4 similar to each other, facial setae formula on article 4 =
3 and 3, slightly oblique to apex, on article 5 = 4 and 4, main spine of article
5 extending to M. 90-100 on article 6, article 5 with no proximoposterior
Spines, spine formula of article 6 = 4 + 4 and 4 + 5 plus middistal seta,
some spines especially long, acclivity on inner margin of dactyls of pereo-
pods 3—4 obsolescent but sharp, emergent setule short, almost fully im-
mersed, midfacial pluseta ordinary. Coxae 5-7 posteroventral setule for-
mula = 1-1-1. Articles 4-5 of pereopods 5-6 narrow, facial spine rows
sparse, facial ridge formula of article 2 on pereopods 5-7 = 0-1-1; width
ratios of articles 2, 4, 5, 6 of pereopod 5 = 51:40:35:13, of pereopod 6 =
69:30:18:9, of pereopod 7 = 80:18:16:8, length ratios of pereopod 5 =
76:34:40:39, of pereopod 6 = 89:51:40:49, of pereopod 7 = 100:21:21:28,
article 2 of pereopod 7 almost reaching middle of article 5, posterior margin
with 10-11 tiny serrations, medial and lateral apices of article 6 deeply
combed, lacking digital processes.

Posteroventral corner of epimeron | rounded, posterior margin weakly
convex, with 2 setule-notches, anteroventral margin with 4—5 short setae,
paired or not, posteroventral face with one medium seta. Posteroventral

VOLUME 94, NUMBER 1 303

corner of epimeron 2 rounded, posterior margin weakly convex, with 2
setule notches, facial setae = 7, posteriormost seta disjunct posteriad. Pos-
teroventral corner of epimeron 3 rounded, weakly protuberant, posterior
margin straight, serrate, setose (2) and with 5 setule notches, ventral margin
with 2 short spine-setae in posterior half.

Urosomite 1 with lateral setule at base of uropod 1, no ventral setae,
articulation line short, urosomite 3 unprotuberant dorsally. Rami of uropods
1—2 with articulate but tightly fixed, deeply immersed and small apical nails,
outer ramus of uropod 1 with one dorsal spine, inner with one, rami of
uropod 2 naked, peduncle of uropod 1 with no basofacial setae and one
small apicolateral spine, medially with 2 marginal spines, apicalmost en-
larged but not displaced. Peduncle of uropod 2 with 4 dorsal spines, medially
with one small apical spine, apicolateral corners of peduncles on uropods
1-2 without comb. Peduncle of uropod 3 with 4 ventral spines, dorsally with
one lateral spine, one medial spine, rami feminine, inner extending to M. 65
on article 1 of outer ramus, apex with one tiny setule, medial and lateral
margins naked, article 2 of outer ramus elongate, 0.40, bearing tiny im-
mersed setule near pointed apex, apicomedial margin of article 1 naked,
lateral margin with 2 acclivities, spine formula = 2-2-2, setal formula = 0.
Telson especially long, length-width ratio = 42:33, almost fully cleft, each
apex of medium width, rounded, scarcely protuberant, lateral acclivities
shallow, narrow, bearing short lateral setule, spine next medial shorter than
setule (thus very short), midlateral setules diverse, largest of medium size.

Male ‘‘d.’’—Ommatidiatarge. Article | of antenna 1 and articles 2—3 of
antenna 2 with medial pubescence; length ratio of articles 1-2 on antenna
1 = 5:3; article 2 with 4 + 1 ventral setae narrowly confined and set api-
cally, primary flagellum with 6 articles, one calceolus each on articles 1-3,
aesthetascs moderately developed, accessory flagellum very short, 4-artic-
ulate. Article 4 of antenna 2 lacking dorsal notch, no dorsal spines and setae,
other setae shortened, facial formula = 3-4-1, article 5 with 3 dorsal sets of
male setae and one calceolus, 3 facial spines, ventrodistal apex with 2 thin
spines; formula of calceoli on flagellum = 1,2,3,5,7,9 ...n. Right lacinia
mobilis broad and scarcely bifid, apices of lobes stubby and rounded, ba-
sofacial setal formula of article 3 on mandibular palp = 0-2.

Setae of coxae 1-3 shortened, coxa 4 not broadened, setae absent. Article
2 of only pereopod 7 narrower than in female. Epimera 1-3 broadened,
posterior margin of epimeron 3 not shortened, setal formulae: epimeron |
anteroventral = 4 (one set paired), posteroventral = 1, posterior = 3 se-
tules, epimeron 2 facial = 6 (posterior one disjunct), posterior = 2 setules,
epimeron 3 posterior = 3 + 3 setules, facial = 0, ventral = 2.

Spine formulae of uropods: uropod 1 peduncle apicolateral = 1, basofa-
cial = | stout and short spine, medial peduncle = only one long apical spine
not displaced, no others; uropod 2 peduncle dorsal = 4, not longer than in

304 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

female, dorsal spines on outer ramus of uropod 1 = 1, of uropod 2 = 0-1,
inner ramus of uropod | = 1, of uropod 2 = 1, ventral spines of peduncle
of uropod 3 = 4, spine formula on article 1 of outer ramus = 2-2-2-2, setal
formula = 0-1-1-1, apex of article 2 with 2 diverse medium setae. Telson
elongate, distal spines not more shortened than already in female, with long
denticle row on each lobe.

Male ‘‘g.’’—Nails on rami of uropods 1-2 enlarged but epimeron 3 with
2 ventral setae, rami of uropod 2 lacking spines, right lacinia mobilis bifid
and outer plate of maxilla 1 with 11 spines. Possible hybrid with spinosus.

Holotype.—Allan Hancock Foundation No. 5746, female “*e’’ 2.77 mm.

Type-locality.—Velero IV 4870, southern California, 33°30'33’N,
117°45'17"W, 10.7 m, benthic orange peel grab.

Voucher material.—Type-locality, male *“‘d’’ 2.71 mm; male “‘g”’ J. L.
Barnard station 38-C-4, Cayucos, California, | July 1961, intertidal, Phyl-
lospadix-pelvetiid community (see J. L. Barnard, 1969a).

This cryptic species is generally similar to E. spinosus but differs in the
following ways: (1) the ommatidia are enlarged; (2) the dorsal notch on
article 4 of antenna 2 has one more seta; (3) the right lacinia mobilis is bifid;
(4) the outer plate of maxilla | has 11 spines; (5) coxa 4 lacks setae; (6)
epimeron 3 has 2 ventral setae; (7) uropods 1-2 lack peduncular comb; (8)
the apical nails of uropods 1-2 are small; (9) uropod | has no basofacial
setae; (10) the rami of uropods 1-2 lack dorsal or marginal spines; (11)
article 2 on the outer ramus of uropod 3 is longer; and (12) has no significant
setae in the female. ;

Just the sparse materials reexamined for this study indicate that charac-
ters 4, 5, 8 and possibly 10 are not very precise and that some hybridization
or gene flow may be occurring between chumashi and spinosus. Neverthe-
less, the species seem to be justified; they appear to be vicariants now
rejoined by human interference, chumashi appearing to be the native Pacific
species and spinosus the Atlantic species that has been brought west.

Distribution.—Southern California, 0-11 m, probably more widespread
(collections of spinosus in Allan Hancock Foundation requiring reidentifi-
cation).

Eobrolgus pontarpioides (Gurjanova), revived
Pararpinia [sic] pontarpioides Gurjanova, 1953:229-230, figs. 11-12.

This species was synonymized with obtusidens by J. L. Barnard —
(1960:249) but is now recognized to be near or in the genus Eobrolgus. It |
is distinguished in the key to Eobrolgus but needs extensive redescription.
No materials are available.

In the original depiction, antenna 1 is drawn as if the setae are apical on —
article 2 but possibly because of the deeply folded condition of article 3 the
true condition of article 2 is obscured. If interpreted as apical, those setae ©

VOLUME 94, NUMBER 1 305

suggest that P. pontarpioides belongs with Eobrolgus, but, if not, the
species probably belongs to Foxiphalus or Grandiphoxus. Gurjanova drew
only one seta on article 3 of antenna 2 suggesting that the species belongs
with Foxiphalus and not with Grandiphoxus where at least 3 setae occur
on article 3. The diversity in head of Foxiphalus and Grandiphoxus is in-
tergraded by Grandiphoxus robustus Gurjanova so that the side view shown
by Gurjanova for P. pontarpioides is useless for.determination. J. L. Bar-
nard (1960) considered P. pontarpioides to be a synonym of Foxiphalus
obtusidens (Alderman) but now that obtusidens has been split into several
species there may be cause to consider P. pontarpioides a good species.
However, epimeron 3 as shown by Gurjanova is not typical of obtusidens
nor of Foxiphalus in general. Pontharpinia pontarpioides must be reserved
as a valid species of unknown genus until further study can be undertaken.

Eyakia J. L. Barnard, 1979

Four species are assigned to this genus; two of these are redescribed in
the system of Barnard and Drummond (1978). These are E. robusta
(Holmes) and E. calcarata (Gurjanova). The other two species, E. uncigera
(Gurjanova) and E. subuncigera (Kudrjaschov) do not occur in Smithsonian
collections.

The two analyzed species are alike in the extra spine on article 4 of
antenna 2, the elongate spine on the molar, the taper on article 2 of pereopod
5, the facial setae on epimeron 3 and the widely spread basofacial setae on
the peduncle of uropod 1.

Key to the Species of Eyakia

[Aes pimneron’s) withlarge tooth 72445. 8s4-25-.-% E. robusta (=?ochotica)
SEE DiImMeronssolacking tOOthos ax WAS VR PL hs es. RO Lice D
Peecicopod / particle 2 iordinany slacking spikem)... S29) 5 E. calcarata
SE ECheOpoda cance a with: langexs pikem. yer nals ites tec d nie te 3
3. Coxa 2 without posteroventral tooth, outer ramus of uropod | with-
OUMMAKeINAl ES PINS Seer PNR aa 3 FOR ARLE, IRA TEER es E. uncigera
— Coxa 2 with posteroventral tooth, outer ramus of uropod 1 with
MAGS IAN LS PATTSS | Kop eeeae, Leng eee awa aise. has Met fs = Le E. subuncigera

Eyakia robusta (Holmes)
Fig. lm, w, y

Paraphoxus robustus Holmes, 1908:518-521, fig. 27.—J. L. Barnard,
1960:235-—236, pl. 25; 1964b:244; 1966a:89; 1966b:29.

Description of female ‘‘w.’’—Head about 21 percent of total body length,
greatest width about 70 percent of length, rostrum unconstricted, broad,
elongate, almost reaching apex of article 2 on antenna |. Eyes small, largely

306 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

occluded with pigment, ommatidia ordinary. Article 1 of peduncle on an-
tenna | about 1.4 times as long as wide, about 2.6 times as wide as article
2, ventral margin with about 11 setules, weakly produced dorsal apex with
2 setules, article 2 about 0.7 times as long as article 1, with midventral row
of 8 setae, primary flagellum with 8 articles, about 0.45 times as long as
peduncle, bearing one short aesthetasc each on articles 2—7, accessory fla-
gellum with 6 articles. Antenna 2 not ensiform, article 3 with thin antero-
dorsal spine and normal facial armament, spine formula on article 4 = 5-5
(tightly compressed) plus offset thin spine midapically, dorsal margin with
notch bearing 5 setae and one spine, ventral margin with 15 groups of 1-3
long to medium setae, one ventrodistal long spine, article 5 about 0.78 times
as long as article 4, facial spine formula = 2-3, dorsal margin bearing one
set of apical setae, ventral margin with 8 sets of one long seta each, 2
ventrodistal long to medium spines set facially; flagellum almost 0.7 times
as long as articles 4—5 of peduncle combined, with 8 articles.

Epistome unproduced. Mandibles with weak palpar hump, right incisor
with 3 teeth, third tooth placed distad, left incisor with 3 humps on 2 branch-
es plus 3—4 weak teeth between, right lacinia mobilis bifid, distal branch
much shorter than proximal, flabellate, tridentate, proximal branch simple,
pointed, with marginal denticles and facial humps, left lacinia mobilis with
4 teeth, middle teeth not shortened, right rakers 11 plus 2 rudimentaries,
left rakers 11 plus one rudimentary, molars composed of short bulbous
humps, each with huge serrate spine, right molar with 2 additional primarily
medium spines, left molar with 3 primarily medium spines, no spine disjunct,
each molar with dense elongate plume; palp article | short, article 2 with 3
long to medium inner apical setae and 4 other long to short inner setae,
article 3 about 1.2 times as long as article 2, oblique apex with 8—10 spine-
setae, basofacial formula = 2-4. Each outer lobe of lower lip with one cone.
Inner plate of maxilla | ordinary, bearing one long apical pluseta, one similar
apicomedial seta, one shorter apicomedial seta and one apicolateral short
seta, outer plate with 11 spines, palp article 2 with one apical spine, 2
apicolateral and 4—5 medial spines and 6 submarginal setae. Inner plate of
maxilla 2 shorter than outer, outer not broader than inner, outer with 5
apicolateral setae, inner with 2 medial setae. Inner plate of maxilliped with
2 large thick apical spines, 2 apicofacial setae, 4 medial setae, outer plate
with 12—14 medial and apical spines plus 3 apical cusps, 4 apicolateral setae,
palp article | with one apicolateral seta, article 2 with one group of 4 api-
colateral setae, medial margin of article 2 moderately setose, article 3 with
4 facial setae, 2 lateral setae, nail of article 4 distinct, short, with 2 accessory
setules.

Coxa | not expanded apically, anterior margin weakly convex, main ven-
tral setae of coxae 1-4 = 23:19:18:29, widely spread, posteriormost seta of
coxae 1-2 shortest, of 3—4 slightly shortened, anterior and posterior margins

VOLUME 94, NUMBER 1 307

of coxa 4 weakly divergent, posterior margin almost straight, posterodorsal
corner rounded, posterodorsal margin ordinary, concave, width-length ra-
tio = 15:18. Gnathopods generally ordinary but hands somewhat slender,
width ratios on articles 5-6 of gnathopods 1-2 = 28:31 and 30:36, length
ratios = 66:71 and 56:73, palmar humps ordinary, palms strongly oblique,
article 5 of gnathopod | elongate, ovate, posterior margin flat, article 5 of
gnathopod 2 ovate, posterior margin triangular.

Pereopods 3-4 similar to each other, facial setae formulas on article 4 =
12 and 12, parallel to apex, on article 5 = 13 and 12, main spine of article
5 extending to M. 100+ on article 6, article 5 with no proximoposterior
spines but with numerous stout setae, spine formula of article 6 = 6 + 4
and 7 + 4 plus tiny middistal seta, one of spines in group of 4 really in
middle, some spines especially long, acclivity on inner margin of dactyls of
pereopods 3—4 absent, emergent setule short, midmarginal pluseta ordinary.
Coxae 5-7 posteroventral seta formula = 11-6-1. Articles 4-5 of pereopods
5-6 narrow, facial spine rows sparse, facial ridge formula of article 2 on
pereopods 5-7 = 0-1-1. Width ratios of articles 2, 4, 5, 6 of pereopod 5 =
42:23:21:11, of pereopod 6 = 69:22:17:10, of pereopod 7 = 84:23:17:8;
length ratios of pereopod 5 = 74:28:35:37, of pereopod 6 = 92:50:60:53, of
pereopod 7 = 100:25:22:33, article 2 of pereopod 7 not reaching middle of
article 4, posterior margin with 6—8 medium serrations, anteroapical margins
of articles 2—3 and anterior margins of articles 4-5 very densely setospinose,
medial apex of article 6 smooth, bearing 6 digital processes; article 2 of
pereopods 5-6 tapering distally, only setulose posteriorly, posteroventral
lobe of pereopod 6 narrow, of pereopod 5 absent.

Posteroventral corner of epimeron | rounded, posterior margin weakly
convex, with 4 widely spread setules, anteroventral margin densely setose,
ventral face with 14+ long setae in irregular ranks (one row of 9 fairly
uniform). Posteroventral corner of epimeron 2 rounded, posterior margin
straight, with 4 widely spread setules, facial setae = horizontal rows of 16,
8 and 4. Posteroventral corner of epimeron 3 with large tooth, posterior
margin straight, with 2 setule notches, ventral margin with 7 setae widely
spread, face with almost horizontal row of 18 setae.

Urosomite | with lateral setule at base of uropod 1 and distoventral setae
besides midventral setae, articulation line complete, urosomite 3 unprotu-
berant dorsally. Rami of uropods 1-2 with articulate but tightly fixed apical
nails, outer ramus of uropod 1 with 3 tightly contiguous dorsal spines, inner
with 2 also tight, outer ramus of uropod 2 with 3 dorsal spines also tight,
inner with no dorsomedial spine, peduncle of uropod 1 with 10 basofacial
setae extending along ventral margin and one apicoventral spine, medially
with 10 thin marginal spines none displaced. Peduncle of uropod 2 with 17
thin dorsal spines, except apicalmost short and stout, medially with one
small thin apical spine, apicolateral corners of peduncles on uropods 1-2

308 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

without comb. Uropod 3 shortened, peduncle with 8 ventral spines, dorsally
with one lateral spine, one medial spine and one setule, rami feminine, inner
extending to M. 50 on article 1 of outer ramus, apex with one seta, medial
and lateral margins naked, article 2 of outer ramus short, 0.13, bearing 2
long setae, medial margin of article 1 with 5 spine-setae, lateral margin with
4 acclivities, short spine formula = 1-1-1-1-2, long spine formula = 0-1-1-1-
1, setae = 0. Telson long, length-width ratio = 15:13, almost fully cleft,
each apex wide, rounded, lateral acclivity broad, shallow, bearing ordinary
lateral setule, thin spine next medial much longer than setule, midlateral
setules diverse or not, largest of medium size.

Female ‘“‘y’’.—Article 2 of antenna | actually with 2 rows of setae, 6
medial long and 8 laterals (3 long, 5 short), flagella with 10 and 7 articles
each. Article 4 of antenna 2 with 10 spines in one row plus one displaced
spine, formula of spines on article 5 = 2 + 2-2, [flagellum broken]. Right
lacinia mobilis with 7 teeth on distal branch; rakers 12 plus 2 rudimentaries;
each molar with 3 spines; basofacial formula on mandibular palp article 3 =
3-3. Inner plate of maxilla 1 normal. Apicolateral setae on outer plate of
maxilla 2 = 7. Outer plate of maxilliped with 5 spines, 3 cusps and 9 lateral
setae.

Setal formula on coxae 1-4 = 29-24-22-34. Epimeron 3 with 10 ventral setae
mostly anterior, main facial row = 13. Uropod 1 peduncle with 3 apicola-
teral spines, inner ramus with only 1. Uropod 2 peduncle with 18 spines,
apical four spines short and stout. Outer ramus of uropod 3 spine formula
lateral = 1-2-2 only, inner ramus only half length of outer ramus article 1,
length of article 2 = 0.07. Lateral setule of telson described for female “*w’’
of spinal dimensions on this specimen.

Another female.—Article | outer ramus of uropod 3 spine formula = 1-2-
2-2-2.

Male ‘‘m.’’—Article 1 of antenna | with medial pubescence, article 2 with
4 lateral setae and ventral groups of 6 and 6. Articles 3—4 of antenna 2 with
medial pubescence, article 4 lacking dorsal notch and spines, lateral formula
on article 4 = 3-6-2, with one spine in group of 3 elongate and apparently
representing displaced spine occurring on female; article 5 with 3 lateral
spines, apex with only 2 spine-setules and one setule, dorsal margin with 8
sets of male setae and probably 7 calceoli [some broken], one flagellar cal-
ceolus each on articles 1,2,3,4,5,6,7,8 . . . n (generic character).

Spines on right mandibular molar = 2, shorter than in female, on left
molar = 3, right lacinia mobilis illustrated.

Pereopods 5-6 like female, pereopod 7 illustrated, with odd article 5 as
shown by Gurjanova (1938, 1951) for calcarata, article 2 narrower than in
female.

Epimera 1-3 broadened, first 2 heavily setose as in female, ventral setae
of epimeron 3 = 8, facial row = 11. Basofacial setae of uropod | = 11,

VOLUME 94, NUMBER | 309

position as in female, but ventral side of peduncle near base with 2-3 large
setae not occurring in female, apicolateral spines of peduncle = 4, outer
ramus = 4, inner = 1. Peduncle of uropod 2 with 25 dorsal spines, basals
shorter than in female, 3 apicalmost short, outer ramus with 6 crowded
spines. Peduncle of uropod 3 with 7 ventral spines, setal formula on article
1 of outer ramus (long and short together) = 1-1-2-2-1-2-1-1-2-2-2-3-3-3-3,
length of article 2 = 0.025!!, with 2 long setae. Telson broadened, length-
width ratio = 8:7, apical elements shortened, composed of one setule and
one spine, each lobe with long row of denticles very densely packed.

Male ‘‘n.’’—Spine formula on article 4 of antenna 2 = 3-7-2.

Voucher material.—Female ‘‘w’’ 9.12 mm, Allan Hancock Foundation
Velero IV 4772, southern California, 80.5 m; male ‘‘m’’ 12.0 mm and female
‘‘y’? 15.0 mm, Albatross, Adakh Island [Alaska], July 1893, surface.

Other material.—Giant female, Port Etches, Alaska, 12—18 fm, 1874, W.
H. Dall 669 (1140), mud; 2 males, Mitrofania Bay, Alaska, Acc. No. 157371,
M.S. [Motorship] Dorothy, light shown over side [of vessel at] 7:30 P.M.,
from surface; giant male, Akun Island, Alaska, surface electric light, A/-
batross, 189?; one male, Old Harbor, Kodiak Island, Alaska, 11 August
1887-1888, Albatross; 2 males, Friday Habor, Washington, USA, 9 August
1950, L. Piternick, light off Oceanographic Lab.

Distribution.—Inner Aleutian Islands southward to Bahia de San Quintin,
Baja California, Mexico, 4—221 m (bottom) and swimming at surface in
neritic zone.

Eyakia calcarata (Gurjanova)

-Parharpinia calcarata Gurjanova, 1938:271—272, 385, figs. 9, 9a, 9b.
Pararpinia [sic] calcarata.—Gurjanova, 1951:388-—392, fig. 237.
Paraphoxus calcaratus.—J. L. Barnard, 1960:238—240, pl. 26; 1966a:88.

Description of female ‘‘n.’’—Head about 20 percent of total body length,
greatest width about 70 percent of length, rostrum unconstricted, broad,
elongate, exceeding middle of article 2 on antenna 1. Eyes medium to large,
clear of pigment (aged in alcohol), ommatidia ordinary. Article 1 of peduncle
on antenna | about 1.3 times as long as wide, about 2.3 times as wide as
article 2, ventral margin with about 9 setules, unproduced dorsal apex with
2 setules, article 2 about 0.6 times as long as article 1, with 2 ventral rows
of 5 and 4 setae, primary flagellum with 12 articles, about as long as pedun-
cle, bearing one short aesthetasc each on articles S—11, accessory flagellum
with 10 articles. Antenna 2 not ensiform, article 3 with one anterodorsal
spine (besides normal facial setules), spine formula on article 4 = 2-4 plus
one long displaced midapical spine, dorsal margin with notch bearing 2 setae
and 2 spines, ventral margin with 6 groups of 2—3 long to medium setae, one
ventrodistal long spine, article 5 about 0.8 times as long as article 4, facial
spine formula = 2-2, dorsal margin bearing one set of small apical setae,

310 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ventral margin with 4 sets of 1-2 long to short setae, 2 ventrodistal long
spines set facially, flagellum proliferating.

Epistome unproduced. Mandibles with weak palpar hump, right incisor
with 3 teeth, third set distad, left incisor with 3 humps on 2 branches and
serrate between, right lacinia mobilis bifid, distal branch much shorter than
proximal, flabellate, quadridentate, proximal branch simple, pointed, with
marginal denticles and facial humps, left lacinia mobilis with 4 teeth plus
One accessory tooth, middle teeth shortened, right rakers 9, left rakers 9
plus 2 rudimentaries, molars composed of bulbous humps, each with huge
serrate spine, right molar with 3 additional medium spines, left with 2 ad-
ditional medium spines, no spines disjunct, each molar with plume; palp
article 1 short, article 2 with one long inner apical seta and 3—4 other long
and short inner setae, article 3 about 1.2 times as long as article 2, oblique
and narrow apex with 8 spine-setae, basofacial formula = 1-1. Each outer
lobe of lower lip with one cone. Inner plate of maxilla | ordinary, bearing
one long apical pluseta, one similar apicomedial seta, 2 apicolateral much
shorter setae, outer plate with 11 spines, palp article 2 with one apical spine,
one lateral and 2 medial spines, 3—4 submarginal setae. Plates of maxilla 2
extending equally, outer not broader than inner, outer with 3 apicolateral
setae, inner with 2 medial setae. Inner plate of maxilliped with 2 large thick
apical spines, 2 apicofacial setae, 5 medial setae, outer plate with 8 medial
and apical spines, 4 apicolateral setae, palp article 1 with one apicolateral
seta, article 2 with one group of 2 apicolateral setae, medial margin of article
2 moderately setose, article 3 with 7 facial setae, 2 lateral setae, nail of
article 4 distinct, short, with 2 accessory setules.

Coxa | expanded apically, anterior margin convex, main ventral setae of
coxae 1-4 = 14-9-9-12, posteriormost seta of coxae 1, 2, 4 shortest, of coxa
3 slightly shortened, anterior and posterior margins of coxa 4 parallel, pos-
terior margin slightly convex, posterodorsal corner sharp-rounded, postero-
dorsal margin ordinary, concave, width-length ratio = 9:10. Gnathopods
generally ordinary but gnathopod 2 significantly stouter than gnathopod 1.
Width ratios on articles 5—6 of gnathopods 1—2 = 28:36 and 28:42, length
ratios = 65:70 and 49:79, palmar humps ordinary, palms strongly oblique,
article 5 of gnathopod | elongate, ovate, posterior margin flat, long, article
5 of gnathopod 2 subtriangular, posterior margin triangular, almost lobate.

Pereopods 3—4 similar to each other, facial setae formula on article 4 =
4 and 5, parallel to apex, on article 5 = 4 and 5, main spine of article 5
extending to M. 85-100 on article 6, article 5 with no proximoposterior
spines, spine formula of article 6 = 4 + 3 and 5 + 3 plus middistal seta, one
of spines in group of 3 situated in middle of posterior articular axis (not in
the normal bilateral fashion), some spines especially long, acclivity on inner
margin of dactyls of pereopods 3—4 absent, emergent setule short, midfacial
pluseta ordinary. Coxae 5—7 posteroventral seta formula = 4-5-1. Articles

VOLUME 94, NUMBER 1 Slit

4_5 of pereopods 5—6 narrow, facial spine rows sparse. Facial ridge formula
of article 2 on pereopods 5-7 = 0-1-1. Width ratios of articles 2, 4, 5, 6 of
pereopod 5 = 45:22:22:13, of pereopod 6 = 75:27:21:11, of pereopod 7 =
88:22:21:11, length ratios of pereopod 5 = 74:28:41:49, of pereopod 6 =
95:58:55:70, of pereopod 7 = 100:25:28:40, article 2 of pereopod 7 reaching
middle of article 4, posterior margin with 8 small serrations, articles mod-
erately setose anteriorly, medial apex of article 6 sparsely and scarcely
combed, bearing 3—4 short digital processes; article 2 of pereopods 5-6
tapering distally, only setulose posteriorly, posteroventral lobe of pereopod
6 broad, of pereopod 5 absent.

Posteroventral corner of epimeron | rounded, weakly protuberant, pos-
terior margin deeply convex, with 3 setule notches, anteroventral margin
with 9 short to medium setae, ventral face with 6 long setae in row running
fully to posterior corner. Posteroventral corner of epimeron 2 rounded,
weakly protuberant, posterior margin barely convex, with 4 setule notches,
facial setae = 9, 2 middle pairs set vertically (setae scattered). Posteroven-
tral corner of epimeron 3 weakly protuberant, with setule sinus, posterior
margin straight, with 2 setule notches, ventral margin with one anterior seta,
face with oblique row of 6 setae.

Urosomite | with lateral setule at base of uropod 1, scarcely any mid-
ventral setae, no ventrodistal setae, articulation line complete, urosomite 3
unprotuberant dorsally. Rami of uropods 1-2 with articulate but tightly fixed
apical nails, outer ramus of uropod | with 3 dorsal spines, not crowded,
inner with 2, outer ramus of uropod 2 with 2 dorsal spines, inner with none,
peduncle of uropod | with 5 basofacial setae extending along ventral margin
and 5 small apicolateral spines, medially with 6 marginal spines, apicalmost
an ordinary spine, none displaced. Peduncle of uropod 2 with 8 dorsal
spines, medially with one small apical spine, apicolateral corners of pedun-
cles on uropods 1-2 without comb. Peduncle of uropod 3 with 4 short ventral
spines, dorsally with one lateral spine, one medial spine and setule, rami
submasculine, inner extending to M. 100+ on article | of outer ramus, apex
with 2 small setae, medial and lateral margins with 7 and one setules, article
2 of outer ramus ordinary, 0.17, bearing 2 medium setae, medial margin of
article 1 with one apical spine and 4—5 medial setules, lateral margin with
3 acclivities, spine formula = 1-1-1-1, short setal formula = 0-1-1-1. Telson
long, length width ratio = 22:19, almost fully cleft, each apex wide, round-
ed, lateral acclivity broad, shallow, with very short lateral and short medial
Spine separated by setule, midlateral setules diverse, larger of medium size.

Voucher material.—Female ‘‘n’’ 6.41 mm, Allan Hancock Foundation
Velero III 171-34, Islas Galapagos, east of Wreck Bay, Chatham Island, 32
fm, 21 January 1934.

Other material.—Velero III 224-34, Colombia, off Gorgona Island, 10 fm,
12 February 1934.

BZ PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Distribution.—Japan Sea, 75 m; California, 324-695 m; Galapagos Is-
lands, 46-73 m; Colombia, Gorgona Island, 18 m.

Eyakia ochotica (Gurjanova)

Pararpinia [sic] ochotica Gurjanova, 1953:225—229, figs. 9, 10.
Paraphoxus ochoticus.—J. L. Barnard, 1960:274.

A possible synonym of E. robusta but see J. L. Barnard (1960).
Distribution.—Okhotsk Sea.

Eyakia subuncigera (Kudrjaschov)
Paraphoxus subuncigerus Kudrjaschov, 1965:1776—-1779, figs. 1, 2.

Article 5 of gnathopod 2 very short.
Distribution.—Okhotsk Sea, 207 m.

Eyakia uncigera (Gurjanova)

Parharpinia uncigera Gurjanova, 1938:267—269, 385, fig. 8.
Pararpinia [sic] uncigera.—Gurjanova, 1951:388, fig. 236.
Paraphoxus uncigerus.—J. L. Barnard, 1960:282.

Distribution.—Japan Sea; Okhotsk Sea.

Literature Cited

Barnard, J. L. 1959. Estuarine Amphipoda in: Ecology of Amphipoda and Polychaeta of

Newport Bay, California.—Allan Hancock Foundation Publications, Occasional Paper

21:13-69, 14 plates.

. 1960. The amphipod family Phoxocephalidae in the eastern Pacific Ocean, with anal-

yses of other species and notes for a revision of the family.—Allan Hancock Pacific

Expeditions 18:175—368, pls. 1-75, 1 chart.

. 1961. Relationship of Californian amphipod faunas in Newport Bay and in the open

sea.—Pacific Naturalist 2:166—186, 2 figs.

. 1964a. Marine Amphipoda of Bahia de San Quintin, Baja California.—Pacific Natu-

ralist 4:55-139, 21 figs., 17 charts, 13 tables.

—. 1964b. Los Anfipodos Bentonicos Marinos de la Costa Occidental de Baja Califor-

nia.—Revista de la Sociedad Mexicana de Historia Natural 14:205—274, 11 figs., 5 tables.

. 1966a. Submarine canyons of southern California Part V Systematics: Amphipoda.—

Allan Hancock Pacific Expeditions 27(5):1—166, figs. 1-46.

. 1966b. Benthic Amphipoda of Monterey Bay, California.—Proceedings of the United

States National Museum 119(3541):1—41, figs. 1-7.

. 1969a. Gammaridean Amphipoda of the rocky intertidal of California: Monterey Bay

to La Jollan—United States National Museum Bulletin 258: 1-230, figs. 1-65.

——. 1969b. A biological survey of Bahia de Los Angeles Gulf of California, Mexico, IV.
Benthic Amphipoda (Crustacea).—Transactions of the San Diego Society of Natural
History 15:175—228, figs. 1-30.

VOLUME 94, NUMBER | S113

—. 1979. Revision of American species of the marine Amphipod Genus Paraphoxus

(Gammaridea: Phoxocephalidae).—Proceedings of the Biological Society of Washington

92:368-379.

, and M. M. Drummond. 1978. Gammaridean Amphipoda of Australia, Part III: The

Phoxocephalidae.—Smithsonian Contributions to Zoology 245:1—-551, figs. 1-269.

Bousfield, E. L. 1973. Shallow-water Gammaridean Amphipoda of New England.—Comstock
Publishing Associates, a division of Cornell University Press, Ithaca & London. vii—xii,
1-312, figs. 1-13, pls. I-LXIX.

Gurjanova, E. F. 1938. Amphipoda, Gammaroidea of Siaukhu Bay and Sudzukhe Bay (Japan

Sea).—Reports of the Japan Sea Hydrobiological Expedition of the Zoological Institute

of the Academy of Sciences, USSR in 1934, 1:241—404, 59 text figs.

. 1951. Bokoplavy morej SSSR 1 sopredel’nykh vod (Amphipoda-Gammaridea).—Aka-

demiia Nauk SSSR, Opredeliteli po Faune SSSR 41:1029 pp., 705 figs.

. 1953. Novye dopolnenia k dal’ nevostochnoi faune morskikh bokoplavov.—Akademlia

Nauk SSSR, Trudy Zoologicheskogo Instituta 13:216—241, 19 figs.

Holmes, S. J. 1905. The Amphipoda of Southern New England.—Bulletin of the United States

Bureau of Fisheries 24:459-529, pls. 1-13, numerous text figs. [unnumbered].

1908. The Amphipoda collected by the U.S. Bureau of Fisheries Steamer, **Alba-
tross,” off the west coast of North America, in 1903 and 1904, with descriptions of a
new family and several new genera and species.—Proceedings of the United States
National Museum 35:489-543, figs. 1-46.

Kudrjaschov, V. A. 1965. Novye vidy bokoplavov (Amphipoda, Gammaridea) iz vostochnoi
chasti Okhotskogo Morja.—Zoologicheski) Zhurnal, Akademiia Nauk SSSR 44:1776—-
1789, figs. 1-10.

Kunkel, B. W. 1918. The Arthrostraca of Connecticut.—Connecticut Geological and Natural
History Survey 6, Bulletin 26 (1) Amphipoda: 15-181, figs. 1-55.

Reish, D. J., and J. L. Barnard. 1967. The benthic Polychaeta and Amphipoda of Morro
Bay, California.—Proceedings of the United States National Museum 120(3565): 1-26.
135 Tle

Shoemaker, C. R. 1925. The Amphipoda collected by the United States Fisheries Steamer
‘‘Albatross” in 1911, chiefly in the Gulf of California.—Bulletin of the American Mu-
seum of Natural History 52:21-61, 26 figs.

Department of Invertebrate Zoology, Smithsonian Institution, Washing-
ton, D.C. 20560.

— an Pau v, oh,
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INFORMATION FOR CONTRIBUTORS

Content.—The Proceedings of the Biological Society of Washington contains papers bearing
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ie CONTENTS

An historical synthesis of Mexican mammalian taxonomy
José Ramirez-Pulido and Marie Claire Britton
Systematics and variation of the Aztec Shiner, Notropis sallei, a cyprinid fish from
Central Mexico Barry Chernoff and Robert Rush Miller
Bernardichthys zorraquinosi, a new genus and species of salmoniform fish from the
late Cretaceous of Oregon Richard W. Huddleston
A revision of the subspecies of Sporophila (‘‘Oryzoborus’’) angolensis (Aves: Em-
berizinae) Storrs L. Olson
Comments on the osteology of balistoid fishes (Tetraodontiformes), with notes on
the triodontid pelvis James C. Tyler and Keiichi Matsuura
A new poison-dart frog (Anura: Dendrobatidae) from the forest of southeastern Colom-
bia William F. Pyburn
On Dorvilleidae and Iphitimidae (Annelida: Polychaeta) with a redescription of Eteonop-
sis geryonicola and a new host record Gary R. Gaston and David A. Benner
A new species of freshwater crab (Crustacea: Anomura: Aeglidae) from insular
south Chile ; Carlos G. Jara and Maria T. Lépez
Taxonomic observations on eastern Pacific Antithamnion species (Rhodophyta:
Ceramiaceae) described by E. Y. Dawson David N. Young
A revision of the northern forms of Euphonia xanthogaster (Aves: Thraupidae)
_ ,Storrs L. Olson
Spadella legazpichessi, a new benthic chaetognath from Enewetak, Marshall
Islands _ Angeles Alvarino
Gorgoniscus incisodactylus, a new isopod of the epicaridean superfamily Cryptonis-
coidea, parasitic on an ascothoracican cirriped from Hawaii Mark J. Grygier
Three new shrimps, and some interesting new records of decapod Crustacea from
a deep-water coral reef in the Florida Keys Robert H. Gore
A new species of Amastigos (Polychaeta: Capitellidae) from the Chesapeake Bay
and Atlantic coast of the Panta States with notes on the Capitellidae of the Chesa-

peake Bay ‘ S70) R. Michael Ewing and Daniel M. Dauer
Uca marguerita, a new species sae fiddler crab aie Ocypodidae) from eastern
Mexico ms $4 Carl L. Thurman II
Additional acanthonotozomatid,. cee yee ng ‘stegocephalid Amphipoda from
the Southern Ocean ci Pane® 3 Les Watling and Heather Holman
A new genus and speciés of Peas Cae Polychaeta) from the north and
south Atlantic 2 Nancy J. Maciolek
Reclassification and redeS¢ription of the, comatulid- Comatonia cristata (Hartlaub)
(Echinodermata: Crinoidea)’ = emalee ATEN Charles Garrett Messing
A new species of Tegastes (Copenods Harpacticoida) associated with a sclerac-
tinian coral at Enewetak Atoll Arthur G. Humes
Three new species of the Eviota epiphanes group having vertical trunk bars (Pisces:
Gobiidae) Susan J. Karnella and Ernest A. Lachner
Taxonomy and postmarsupial development of a dominant deep-sea eurycopid
isopod (Crustacea) George D. Wilson

The amphipod genera Eobrolgus and Eyakia (Crustacea: Phoxocephalidae) in the
Pacific Ocean J. Laurens Barnard and Charline M. Barnard

101

107

122

135

163

169

181

228

240

254

264

276

295

74.0615

(ISSN 0006-324X)

Proceedings
of the
BIOLOGICAL SOCIETY
of
WASHINGTON

Volume 94 20 July 1981 Number 2

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

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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Washington. Correspondence concerning membership should be addressed to the Treasurer,
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tution, Washington, D.C. 20560.

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Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044

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

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 315-343

THE TANAIDAE (CRUSTACEA; TANAIDACEA) OF
CALIFORNIA, WITH A KEY TO
THE WORLD GENERA

Jurgen Sieg and Richard N. Winn

Abstract.—Identification keys to the world genera and the California
species of the family Tanaidae are presented. Diagnostic characters are
discussed for Sinelobus stanfordi (Richardson), Pancolus californiensis
Richardson, Zeuxo normani (Richardson), and Zeuxo paranormani Sieg.
Two new species, Synaptotanais notabilis and Anatanais pseudonormani
are described. Three additional species, Zeuxo coralensis Sieg, Zeuxo ma-
ledivensis Sieg, and Zeuxo seurati (Nobili), known to occur within the east-
ern Pacific region, are included in the identification keys. The distribution
of all species are discussed.

Investigations in recent years have indicated that the tanaidacean Crus-
tacea are among the more numerically abundant invertebrates of certain
marine habitats (e.g. Livingston, 1977; Odum and Heald, 1972; Gage and
Coghill, 1977). However, difficulties in identifying specimens to species or
even family have impeded research on their biology. The identification keys
to the suborders and families of the Tanaidacea by Sieg and Winn (1978)
were an outcome of an examination of tanaids from benthic macrofaunal
samples from the southern California borderland. It was stated then that
additional identification keys, new species descriptions, and discussions of
the families occurring along the California coast would be subsequently
presented. The following is an examination of the members of the family
Tanaidae from California.

Tanaidae Dana, 1849

Little is reported on the members of the family Tanaidae from California.
Tanais normani Richardson and Pancolus californiensis Richardson, were
originally described from Monterey Bay in northern California. Localities
in southern California were noted for ‘‘Anatanais normani’’ by Miller (1968)
at Catalina Island, Anacapa Island, Santa Barbara Harbor, and Moss Beach.
Dillon Beach was mentioned as another locality for Pancolus californiensis
by Lang (1961).

Miller (1968), and Miller and Menzies (1954) reported a Tanais sp., which
was shown to be a male of Sinelobus stanfordi (Richardson) by Sieg (1980),

316 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

who also showed that the type-material of Tanais normani Richardson ac-
tually consisted of 2 species of Zeuxo, one of which is now referred to
Zeuxo paranormani Sieg (1979).

None of these previously known species was collected from the benthic
sampling program (Bureau of Land Management 1976-1977) off southern
California, which emphasized shallow subtidal regions (13 m) to deep shelf,
slope, and basin soft-bottom regions (1886 m). Two new species within the
family Tanaidae, Synaptotanais notabilis and Anatanais pseudonormani,
were found and are described here.

Several shallow, near-shore studies recently completed or now in progress
have indicated the frequent presence of tanaidaceans in great numbers, up
to thousands per m? (Muscat, pers. comm.; Hammer and Zimmerman, 1979;
Straughan 1977). The highest density with 140,000/m? has been reported for
Kalliapseudes crassus by Barnard (1970).

There are currently 6 known species of Tanaidae from California: Ana-
tanais pseudonormani n. sp.; Pancolus californiensis Richardson, Sinelo-
bus stanfordi (Richardson), Synaptotanais notabilis n. sp., Zeuxo normani
(Richardson), and Zeuxo paranormani Sieg (for futher details see Sieg,
1980). Three additional species known from the eastern Pacific region,
Zeuxo maledivensis Sieg, Z. seurati (Nobili), and Z. coralensis Sieg are
included in the following key to the California species of Tanaidae, since
there is some possibility they may be found there also.

The terminology used is based on that suggested by Racovitza (1923) for
the Isopoda, with modifications proposed by Sieg (1973, 1977, 1980), which
includes a discussion of setae and spines of the Tanaidacea.

There has been much variation in the nomenclature applied to body seg-
ments and appendages for the Tanaidacea as discussed by Hansen (1913),
Lang (1953), Wolff (1956, 1962) and Gardiner (1975). Gardiner summarized
Wolff’s terminology and suggested alternative terms to those used by Lang
(1968). Lang’s system is adopted here for the orientation of the appendages
where: dorsal, ventral, medial, lateral are equivalent to tergal, sternal, ros-
tral, and caudal, referring to limbs directed ventrally and laterally from the
body, in their natural position. The abbreviations used for the figures are:
A.1 (Antenna 1), A.2 (Antenna 2), L (Labrum), Md, (left mandible), Md,
(right mandible), La (labium), Mx.1 (maxilla 1) Mx.2 (maxilla 2), Epi (epig-
nath), Mxp (maxilliped), che (cheliped), P.1—P.6 (pereopod 1-6), Pl. 1-5
(pleopod 1-5), Plt (pleotelson) and Uro (uropod).

Key to the World Genera of the Family Tanaidae .
(Modified from Sieg, 1980)
(*Not yet recorded from California)

1. Uropods long and slender, articles more than twice as long as broad
CBige PUTO! Tyo os Ve oes ec forieca sags Sosa shes eee eee y)

VOLUME 94, NUMBER 2 7)

Uro:]

Uro:4

A.l:1 Al:2

Fig. 1. Uropods and antenna | of different members of Tanaidae. Uro: 1—uropod long and
slender, articles more than twice as long as broad; Uro: 2—uropod short, articles twice as long
as broad; Uro: 3—uropod with terminal article normal; Uro: 4—uropod with terminal article
greatly reduced; A.1: 1—antenna 1 with first article twice length of second article; A.1: 2—
antenna 1 with first article more than 2.5 times length of second article.

— Uropods short, articles twice (rarely 3 times) as long as broad (Fig.
1 LOWROS 2A ea pa ho, estrone alee Wei SOR LUN viNes ttre ey aes gn Re a DP 5

2. Antenna 1 of 5 articles, first and second article with many distal
setae; cephalothorax broader than long ........ Archaeotanais Sieg*

318

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

oes

Fig. 2. Synaptotanais notabilis,female.

VOLUME 94, NUMBER 2

05mm

Fig. 3. Synaptotanais notabilis, female.

319

320

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Antenna 1 of 4 articles, first and second article with few distal
setae; cephalothorax longer than broad (Uro about ‘‘6 to 11-
goimted’’)..5 PY: hake 05 eka Cees Uae. eS SP Nec eR ee 3

. Endopod of uropods of at least 10 articles; antenna 2 of 8 articles;

outer lobe of labium with spine ................. Langitanais Sieg*
Endopod of uropods of 5—6 articles; antenna 2 of 7 articles; outer
lobe of labium without spine, with or without terminal article (Uro

* 6=777ommted ) cee vw bw si TP oa al ee 4
. Outer lobe of labium without terminal article; tergite of fifth pleon-
ite not distinctly separate from pleotelson ......... Protanais Sieg*
Outer lobe of labium with terminal article; tergite of fifth pleonite
distinctly separate from pleotelson ............. Synaptotanais Sieg
. Terminal article of uropods greatly reduced (Fig. 1, Uro: 4) ..... 6
Terminal article of uropods normal, not reduced (Fig. 1, Uro:
SES Se eietae Ser, em teanae ian any okt Meme So 5 5 6 5 5 00 o 8
. Pleopod 3 reduced; carpal spines of pereopods 2—4 strongly pro-
GU CE EE | 0 ls ie NS oi eo al ape ee = “ay suelel oss a eee 7
Pleopod 3 normal, not reduced; carpal spines of pereopods 2-4
normal, not strongly produced.................... Zeuxoides Sieg*
. Antenna 1 of 3 articles; pleon with 3 pleonites (tergites) visible
CORSET ASAD sick. ig ek ees. athe al a tbe etanene wrod eles 2 Pancolus Richardson
Antenna 1 of 4 articles; pleon with 5 pleonites (tergites) visible
dorsally) plusepleotelson 4. - 4.64 - aoe ee ee Pancoloides Sieg
. Pleon with 4 pleonites (tergites) visible dorsally, plus pleotelson . 11
Pleon with 5 pleonites (tergites) visible dorsally, plus pleotelson.. 9

. Coxa of pereopod | without protuberance; pereonites 1-3 com-

bined not longer than broad; terminal lobe of outer lobe of labium
not well separated; chelae of male greatly enlarged, therefore,
cephalothorax of males in cross-section nearly triangular

LE SO ae Oe ane eM ore orn cera) Stan Somer th Hexapleomera Dudich*
Coxa of pereopod | with or without protuberance; pereonites 1-3
large, combined longer than broad; terminal lobe of outer lobe of
labium well separated; chelae of male not greatly enlarged, there-

fore, cephalothorax of male normal ......................20--- 10
. First article of antenna 1 greater than 2.5 times, often 3 times,

length of second article (Fig. 1, A.1: 2) ........... Zeuxo Templeton

First article of antenna | twice length of second article (Fig. 1, A.1:

BY Sea hcaahe oro eae ea ede vie en ee ee Anatanais Nordenstam
s Wropods..Of 7 articles: ci. saves Aes ae eee Arctotanais Sieg

Uropods’ of 2=5 articles .2... 292 22k). Ge Re eee 12

. Pleonites | and 2 with strong transverse rows of long plumose

NDEI See She a Si Rete en eed ase ne eee 13

VOLUME 94, NUMBER 2 321

Fig. 4. Synaptotanais notabilis, female.

322

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Pleonites 1 and 2 without transverse rows of setae, but with a few

lateralplumose hairs” 2. see oetaere a Monoditanais Sieg*
. Outer lobe of labium with terminal lobe .......... Tanais Latreille*
Outer lobe of labium without terminal lobe .................... 14

. Fourth article of antenna 2 with circle of setae .................

Pe EC Re ee ROLE, ae ve on RCE gener ten ae Parasinelobus Sieg*
Fourth article of antenna 2 without circle of setae, but with few
GISTAIGSE TAC cu LR Oe CS i eos LS on a NSN Sinelobus Sieg

Key to the Species of California Tanaidae

. Uropods long and slender, articles more than twice as long as broad

Gicalh.Unom I rics, 2-55.12) aos Synaptotanais notabilis n. sp.
Uropods short, articles twice as long as broad (rarely 3 times) (Fig.
RU TOL DQ) a Ses a us I RR A a, 8 2

. Pleon of 3 pleonites plus pleotelson; pleopod 3 reduced (Fig. 7)

PPR An We Nina ce LM feat ts Pancolus californiensis Richardson
Pleon of 4—5 pleonites plus pleotelson; pleopod 3 not reduced..... 3

. Pleon of 4 pleonites plus pleotelson; pleonites 1 and 2 each with well

developed transverse row of long plumose setae (Fig. 6) .........
OMNI le Sis sre IMI cc uence Sinelobus stanfordi (Richardson)
Pleon of 5 pleonites plus pleotelson; pleonites 1 and 2 with few

lateral, plumoOSe ‘SCtae ney. so. ee ek oe oe ee 4
. First article of antenna 1 2.0 times length of second article (Fig. 1,

Feo a |) Meena a Sal SONS Prem Pooper es Serennery er Anatanais pseudonormani 0. sp.

First article of antenna | 2.5 times length of second article, nearly

37.0: times (Eig. 5 Ale ds: 2) rcssnectire bcp syeyteyssickolerstaus is i = cute vag eRe ene 5
. Uropods of 4 articles (adults); coxa of pereopod 1 produced slightly

(CE51@ sil 5) 8 Bisnstichas'susge rcv ie eee ee ee kegs @ SE Zeuxo coralensis Sieg*

Uropods of 5=6-articles (adults) oo ect. Ove Bs ole 6

. Lacina mobilis of right mandible reduced to small projection (Fig.

17); coxa of pereopod 1 produced slightly .... Zeuxo seurati (Nobili)*
Lacina mobilis of right mandible only slightly reduced (Figs. 13-16)

. Coxa of pereopod 1 with strong, triangular protuberance (Figs. 13,

14); inner border of endopod of pleopods with more than 1| proximal
SOLA) cduced.s auhviwinss caw dae eRe Beem arene ee 8
Coxa of pereopod | with small, triangular protuberance (Fig. 16);
endopod of pleopods with | proximal seta.. Zeuxo maledivensis Sieg*

. Uropods of 6 articles (adults, Fig. 13); coxa of pereopod 1 with large

protuberance (Fig. 13); carpus of pereopod 2 with 4 caudal and 2
rostral spines (Fig. 13): bases. eee Zeuxo normani (Richardson)
Uropods of 5 articles (adults, Fig. 14); coxa of pereopod | with less

VOLUME 94, NUMBER 2

Fig. 5. Synaptotanais notabilis, female.

pronounced protuberance: carpus of pereopod 2 with 3 caudal and
PLOSPLALSPINE SW OAIe, Lome eee Gene ER). Zeuxo paranormani Sieg

Synaptotanais notabilis, new species
Figs. 2-5, 12

Description (females, neuters).—Body length of fully developed speci-

mens about 7 mm.; manca stages and neuters smaller; slightly more than 4
times as long as broad.

Cephalothorax smoothly rounded; narrowing from posterior to anterior;
1.2 times longer than broad, with small rostral eye-lobes separated from

323

324 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON _

cephalothorax by concave sutures; | small seta nearby, and additional lat-
eral seta.

Pereon with first 3 pereonites laterally rounded in dorsal view. Last 3
pereonites with swollen region posteriorly. Pereonite 1 3.5 times broader
than long; anterior border smoothly concave, with few setae anterolaterally
near exhalent opening. Pereonite 2—3 times broader than long, lateral margin
with | seta rostrally and | at the middle. Pereonite 3 twice as broad as long,
bearing setae similar to pereonite 2. Pereonite 4 1.2 times broader than long,
with | seta in each lateral corner. Pereonite 6 1.5 times broader than long,
similar to pereonites 4 and 5.

Pleon with 5 distinct segments plus pleotelson visible in dorsal view. Last
2 segments smaller. Fifth pleonite (sternite) fused ventrally with pleotelson.
Pleonites 1-3 with lateral plumose setae.

Antenna 1 (Fig. 2) of 4 articles. First article relatively large, 5.0 times
longer than broad; inner border with 3 plumose hairs proximally, 5 setae
and 2 plumose hairs distally; outer border with 1 seta and 1 plumose hair
distally. Second article twice as long as broad; inner border with 4 distal
setae; outer border with 3 distal setae. Third article small, but also twice as
long as broad; inner border and outer border each with | distal seta. Fourth
article minute, conical, with 3 short setae, 7 long setae and 3 esthetascs.

Antenna 2 (Fig. 2) of 7 articles. First article fused with cephalothorax, as
long as broad, without setae. Second article twice as long as broad; outer
border with flange, with distal seta; inner border with 2 setae. Third article
small, as long as broad, without setae. Fourth article 4.0 times longer than
broad; inner border with 3 distal setae; outer border with 1 seta and 2
plumose hair. Fifth article 2.5 times longer than broad; inner border with 2
setae and | plumose hair distally; outer border with 1 seta and 1 plumose
hair. Sixth article as long as broad; inner and outer border with | distal seta.
Seventh article minute, peak-like, with 4 short and 4 long setae.

Labrum (Fig. 3) hood-like, completely covered with fine hairs.

Mandible (Fig. 3) strongly produced. 0.33 of border of the terminal face
of the pars molaris toothed. Lacinia mobilis of left mandible well developed,
border distally crenulate, with 2 plumose membranous spines near articu-
lation. Lacinia mobilis of right mandible distinctly smaller than left.

Labium (Fig. 3) with an inner and outer lobe, divided deeply in middle,
distally covered with fine hairs. Outer lobe with terminal article and proxi-
mally with small lateral spines.

Maxilla | (Fig. 3) with uniarticulate palp bearing 4 fine, plumose terminal
setae; endite with 8 spines surrounded by distal setae.

Maxilla 2 (Fig. 3) long, oval, with small terminal setae.

Maxilliped (Fig. 5) developed normally; coxa perpendicular to basis, with
2 mid-ventral setae; not fused medially. Basis not fused medially; also with
1 mid-dorsal seta. Palp of exopod of 4 articles. First article 1.5 times longer

VOLUME 94, NUMBER 2 B25

than broad, with 2 setae on outer border. Second article triangular; outer
border concave, bearing | long seta; inner border bearing 3 plumose and 6
naked setae. Third article twice as long as broad; inner border slightly con-
vex, with 2 rows of 8 and 4 setae. Fourth article 3.0 times longer than broad;
outer border with 1 distal seta and inner border with 2 rows of 6 setae each.
Endopod (inner lobe) not fused medially; with 2 oblique setae distally and
2 membranous spines medially (‘‘coupling-hooks’’sensu Lang, 1968).

Epignath (Fig. 3) kidney-shaped, with small appendix, completely cov-
ered with fine hairs.

Cheliped (Fig. 2) well developed, but slender. Basis twice as long as
broad, with medial seta mid-ventrally. Merus triangular; ventrally with 3
medial setae. Carpus small, about twice as long as broad; ventrally with 3
medial setae on the distal third; mid-dorsally with 1 seta and distally with
3 medial setae. Propodus and fixed finger slightly shorter than carpus, with
1 medial and | lateral seta near articulation of dactylus. Fixed finger with
Spine at its tip and 2 lateral setae; dorsal border with 7, and ventral border
with 5 setae. Dactylus with spine, curved slightly.

Pereopod | (Fig. 4) long and slender. Coxa having non-moveable articu-
lation with sternite, with long prominent protuberance bearing 2 setae. Basis
more than 6.0 times longer than broad; proximal third with 2 small sternal
setae; distally with 1 tergal seta. Ischium absent. Merus 3.5 times longer
than broad; sternally and tergally each with caudal seta distally. Carpus
about 5.0 times longer than broad; tergally with 2 and sternally with 1 lateral
setae distally. Propodus slender, more than 7.0 times longer than broad;
middle with rostral seta sternally. Dactylus and spine combined slightly
longer than 0.5 length of propodus.

Pereopod 2 (Fig. 4) with coxa fused with sternite, without protuberance.
Basis 4.5 times as long as broad; sternally with | proximal seta; tergally
with 1 distal seta. Ischium absent. Merus 3.5 times longer than broad; ter-
gally with 3 caudal setae distally and | caudal spine; sternally with caudal
seta. Carpus twice as long as broad; with 3 caudal setae distally and nearby
medial spine sternally; distal tergal border with 3 caudal spines, 2 caudal
setae and 5 rostral spines, 2 of which are very small. Propodus 4.5 times as
long as broad; tergal row of 4 caudal setae and rostral seta distally; sternally
with 1 caudal and 1 rostral seta distally. Dactylus and spine combined 0.5
as long as propodus.

Pereopod 3 (Fig. 4) proportions and armament mainly similar to pereopod
2. Carpus tergally with 4 caudal spines and caudal seta distally. Propodus
tergally with 1 rostral spine.

Pereopod 4 (Fig. 4) coxa fused with sternite, with no distinguishing fea-
tures. Basis 4.0 times longer than broad; middle and distal portions each
with 3 tergal setae. Ischium absent. Merus 2.5 times longer than broad;
distally with caudal and rostral seta sternally, and caudal and rostral spine

326 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tergally surrounding 2 setae. Carpus twice as long as broad, with ventral
indenture for the propodus and 3 caudal and 2 rostral setae; distal row of
dorsal spines consists of 6 caudal and rostral spines each. Propodus 3.5
times longer than broad; sternally with plumose hair and | rostral and 1
caudal seta; tergally with 2 setae near midlength and distally with 1 rostral
and 1 caudal seta. Dactylus and spine coalesced to claw bearing row of
membranous spines rostrally and caudally.

Pereopod 5 (Fig. 4) proportions and armament as in pereopod 4. Carpus
sternally with 2 rostral and 2 caudal distal setae; tergally with row of 7
spines rostrally and caudally.

Pereopod 6 (Fig. 4) proportions and armament similar to 4 and 5. Carpus
with 6 caudal and rostral spines distally. Propodus bears distal row of scal-
pelliform setae tergally; sternally with 7 plumose setae and plumose hair
between long caudal and rostral setae.

Pleopods (Fig. 2) similar, not reduced. Outer border of basis with 5 setae,
inner border with 1 seta. Exopod of 1 article, without setae on inner border
and with many setae on outer border. Endopod of | article, with some setae
on proximal inner border; outer border bears many setae, distal seta bicil-
lated (specialized).

Pleotelson (Fig. 5) caudal point prominent, with 2 long setae; 1.5 times
broader than long, with 2 long and 2 shorter caudal setae near articulation
with uropods. Last pleonite partly fused with pleotelson; tergally well
marked, sternally totally fused.

Uropods (Fig. 5) long and slender; consisting of basis and 6-articled en-
dopod (Uro: ‘‘7-jointed’’). Basis 5.0 times as long as broad, distally with 3
long and 2 short setae. Endopod with first article less than 4.0 times as long
as broad, distally with 2 short setae. Second and third article less than 3.0
times as long as broad; second with 3 distal, third with 2 distal setae and a
plumose hair. Fourth and fifth article 5.0 time as long as broad; fourth with
3 short setae and | plumose hair distally. Sixth article 4.0 times as long as
broad, with 4 long and 2 short setae, and 2 plumose hairs distally.

Etymology.—The specific name is taken from the Latin word for ‘‘nota-
ble.”’

Material.—Velero IV sta. 23092/BLM 182 (33°39'0"N, 120°6'1"W), off
Santa Rosa Island, California, 133 m: 5 2 9:1 ex. Senckenberg Museum
Frankfurt SMF 8674, 2 sp. coll. Sieg, 2 ex. USNM 170654. Velero IV sta.
23093/BLM 183 (33°39’0"N, 119°58’0’W), off Santa Rosa Island, 133 m: 1
2, University of Southern California, Allan Hancock Foundation. Velero
IV sta. 23187/BLM 79 (33°45'9’"N, 120°03’0"W), off Santa Rosa Island, 133
m: 2 292, University of Southern California, Allan Hancock Foundation.

All specimens are syntypes, with Velero IV sta. 23092 designated as type-
locality.

Discussion.—This species is readily distinguishable by its slender uropods

VOLUME 94, NUMBER 2

A
|

|
on

|
<

38

We

N

Imm

is ae)

yes AUS Fw
I aaa 4 sa Ae
y fp eX
\ dr (aay

UO

ASFA

Fig. 6. Sinelobus stanfordi, female and male.

327.

328 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Lmm

Fig. 7. Pancolus californiensis, female.

VOLUME 94, NUMBER 2 329

and relatively long first articles of antenna 1. Its present distribution is
limited to the southern California borderland.

Sinelobus stanfordi (Richardson, 1901)
Fig. 6

Synonymy, see Sieg (1980).

Discussion.—The ‘‘Tanais’’ sp. mentioned by Miller (1968), and Miller
and Menzies (1954) belongs to Sinelobus stanfordi as noted by Sieg (1980).
The species may be recognized by the 5-segmented pleon (4 pleonites and
the pleotelson), first and second pleonite with 2 curved rows of long, plu-
mose vertical setae. The third pleonite also bears some plumose setae. Si-
nelobus is distinguished from the “‘true’’ Tanais by the loss of the terminal
lobe in the labium and by the cephalothorax of males narrowing markedly
from posterior to anterior (Fig. 6). The species is cosmopolitan and occurs
in shallow intertidal and estuarine areas, including some records from fresh
water.

Pancolus californiensis Richardson, 1905
Figes

Synonymy, see Sieg (1980).

Discussion.—This species is easily recognized by its 4-segmented pleon
(3 pleonites and the pleotelson). There appears to be no other species which
share this character. Good descriptions and discussions of affinities have
been given by Lang (1960) and Sieg (1980). The known distribution is from
Monterey Bay in northern California (Richardson, 1905) to central California
(although few records are available), and into southern California. In recent
studies within southern California, Pancolus californiensis has been found
mainly in intertidal and shallow subtidal areas associated with algae, or
coarse-grained, sandy beaches.

Anatanais pseudonormani, new species
Figs. 8—11, 12

Description. (females, neuters).—Length of body 4-5 mm. Lateral margin
of cephalothorax not rounded, slightly narrowed from posterior to anterior,
1.2 times longer than broad, with small rostral indentations separating eye-
lobes, with 2 small setae nearby and an additional lateral seta.

Pereonites similarily rounded laterally in dorsal view, with seta in antero-
lateral corners and occasionally in middle. Pereonite 1 4.5 times as broad
as long; anterior border smoothly concave, with some setae near exhalent
opening. Pereonite 2 3.0 times, and pereonite 3 2.3 times broader than long.
Following 3 segments broaden posteriorly. Pereonites 4 and 5 1.8 times
broader than long. Pereonite 6 twice as broad as long.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

330

Fig. 8. Anatanais pseudonormani, female.

VOLUME 94, NUMBER 2 331

Fig. 9. Anatanais pseudonormani, female.

Pleon (Fig. 8) with 5 segments visible in dorsal view, plus pleotelson.
First 3 pleonites relatively large, posterolaterally produced, bearing some
plumose setae. Pereonites 1-3 (in dorsal view) 5.0 times broader than long;
4 and 5 small and compressed anterior to posteriorly; nearly 7.0 times broad-
er than long, with 2 lateral setae.

Antenna | (Fig. 8) of 4 articles. First article 4.0 times longer than broad;
proximal 0.33 of outer border with 2 plumose setae; outer border with 4
setae and inner border with 4 plumose setae and 1 normal seta distally.

B82 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Second article nearly 3.0 times longer than broad; outer border with 3 setae
and 5 plumose setae distally; inner border with 4 distal setae. Third article
also 3.0 times longer than broad; outer border with 2 and inner border with
3 setae distally. Fourth article minute, conical with 2 esthetascs, 2 plumose,
2 short, 2 very long and 4 other setae.

Antenna 2 (Fig. 8) of 7 articles. First article fused with cephalothorax; as
long as broad, without setae. Second article 1.5 times longer than broad;
inner border with 1 seta proximally and distally. Third article also small, as
long as broad, without setae. Fourth article 2.7 times longer than broad;
inner border with 4 setae; outer border with 3 normal and 3 plumose setae
distally. Sixth article somewhat longer than broad, with 3 setae. Seventh
article minute, conical, with 1 short and 5 long setae.

Labrum (Fig. 9) hood-shaped, completely covered with fine hairs.

Mandibles (Fig. 9) developed normally; 0.33 of border of terminal face of
pars molaris toothed. Lacinia mobilis of left mandible well developed and
distally with round teeth, with 2 plumose membranous spines near its artic-
ulation. Lacinia mobilis of right mandible reduced, but recognizable, bearing
2 plumose membranous spines near its articulation.

Labium (Fig. 9) with inner and outer lobes, both deeply incised in middle.
Distal part of inner lobe with fine hairs. Outer lobe with terminal article,
distally covered with fine hairs and row of small lateral spines proximally.

Maxilla 1 (Fig. 9) with endite and uniarticulate palp. Endite with 8 stout
terminal spines surrounded by circle of setae. Palp as long as endite, with
5 very fine plumose setae.

Maxilla 2 (Fig. 9) long, oval with 2 small setae.

Maxilliped (Fig. 8) developed normally. Coxa perpendicular to basis, not
fused medially, with 1 mid-ventral seta. Basis slender and small when com-
pared with palp; twice as long as broad, not fused medially, with only 1
small seta near articulation of palp. Palp of 4 articles; first article with outer
border convex, 1.5 times longer than broad, with 1 seta. Second article
triangular; outer border concave, bearing | seta; inner border with 3 strong
plumose setae, and 7 normal setae. Third article 2.0 times longer than broad,
inner border slightly convex with 2 rows of 7 setae each. Fourth article
nearly 4.0 times as long as broad; outer border with 1 distal seta; inner
border with 2 rows of 7 setae. Endopod (inner lobe) not fused, distally
covered with fine hairs and with 2 biciliated (specialized) setae on each side,
as well as 3 small membranous projections (“‘coupling hooks,”’ sensu Lang,
1968).

Epignath (Fig. 9) kidney-shaped, with small appendix, completely cov-
ered with fine hairs.

Cheliped (Fig. 11) slender; coxa behind proximal conjunction of basis,
latter article more than twice as long as broad, distally with a rostral seta
sternally. Merus triangular; distally with 2 rostral setae sternally. Carpus

VOLUME 94, NUMBER 2

Fig. 10. Anatanais pseudonormani, female.

333

334

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 11.

Anatanais pseudonormani, female.

VOLUME 94, NUMBER 2 335

sc. SANTA BARBARA

SOUTHERN CALIFORNIA
BORDERLAND

Rosa_—~___-_~ Santa

Is Cruz Is an fee a : ee aaa fy
=> L SR OORED ‘

¢_SAN DIEGO
i Bank ae Wh:
° \ 2 eal
e A.pseudonormani sp.n
4 S.notabilis sp.n.

Fig. 12. Known distribution of Synaptotanais notabilis and Anatanais pseudonormani in
the southern California borderland.

twice as long as broad, with 1 seta mid-tergally and 2 distal setae; sternally
distal third bears group of 5 setae, inserting variously caudally or sternally.
Propodus and fixed finger slender, as long as carpus, with 2 rostral setae
near articulation of dactylus distally. Fixed finger with spine at tip and 2
caudal setae; dorsal border with 5 rostral and sternal border with 4 rostral
setae. Dactylus with spine, slightly curved.

Pereopod 1 (Fig. 10) coxa fused with sternite, with prominent protuber-
ance bearing 3 setae. Basis 5.5 times longer than broad, proximal third with
1 seta sternally, with another tergal seta distally. Ischium absent. Merus 1.5
times longer than broad; distally with 1 rostral seta tergally and sternally.
Carpus twice as long as broad; distally with 1 caudal seta tergally and ster-
nally. Propodus 4.7 times longer than broad, distal third with 4 tergal-caudal
setae, some plumose hairs, and | caudal seta distal-sternally. Dactylus and
Spine combined as long as propodus.

Pereopod 2 (Fig. 10) with coxa fused to sternite, without specialized fea-
tures. Basis 3.5 times longer than broad, proximal third with plumose hair

336 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

distally and dorsal seta distally. Ischium absent. Merus twice as long as
broad, with tergal-caudal spine, | caudal and 1 rostral seta distally. Carpus
1.5 times longer than broad; sternally with 1 caudal seta distally; tergally
with 1 spine and 1 seta caudally and 3 rostral spines, first twice length of
second. Propodus 3.5 times longer than broad, distal third with 2 setae
tergally and 1 seta sternally. Dactylus and spine combined slightly longer
than 0.66 length of the propodus.

Pereopod 3 (Fig. 10) proportions and armament similar to pereopod 2.

Pereopod 4 (Fig. 10) strongly produced. Coxa fused to sternite, with no
characteristic features. Basis 2.5 times longer than broad; proximal half with
2 plumose hairs sternally; tergally with 2 feathered hairs and | seta. Ischium
absent. Merus twice as long as broad, slightly curved sternally, with 1 caudal
seta; distally with 1 caudal and | rostral spine surrounding | seta tergally.
Carpus twice as long as broad; sternally with distal indentation for propodus
and | caudal and rostral seta distally. Caudal and rostral distal rows of
spines consist of 4 spines each. Propodus 2.7 times as long as broad; tergally
with | seta mid-tergally as well as a small caudal and rostral seta distally;
sternally with a plumose hair, 1 long caudal and 1 rostral seta distally.
Dactylus and spine fused to claw bearing row of membranous spines ros-
trally and caudally.

Pereopod 5 (Fig. 10) proportions and armament similar to pereopod 4.

Pereopod 6 (Fig. 10) proportions and armament similar to pereopod 4 and
5, but with a distal row of only 4 caudal spines dorsally. Propodus with
additional short row of scalpelliform setae distally: sternally with small seta
Setween long caudal and rostral setae.

Pleopods (Fig. 11) similar. Outer border of basis with 3 setae, inner border
with 1 seta. Exopod of 1 article; outer border bearing many setae. Endopod
of 1 article; inner border with 2 plumose proximal setae, outer border with
many plumose setae, most distal one oblique (specialized).

Pleotelson (Fig. 11) 1.5 times broader than long, caudal point prominent,
with 2 long setae. Caudal border with 2 additional setae on each side of
caudal point, | seta laterally. Last 2 pleonites fused with pleotelson; tergites
small but well marked; sternites partly fused with pleotelson.

Uropods (Fig. 11) with basis (Uro: **6-jointed’’), endopod of 5 articles.
Basis 2.5 times longer than broad; outer border with 3 distal and inner
border with 1 distal seta. First 3 articles 1.4 times longer than broad. First
article without distal setae. Second article with 2 setae along inner border.
Third article with 2 setae along outer border and 3 setae along inner border.
Fourth article 2.5 times longer than broad, with 4 distal setae along outer
border and 5 distal setae along inner border. Fifth article twice as long as
broad, with 6 long and 2 short setae.

Etymology.—The name was chosen because of the close resemblance to
Zeuxo normani (Richardson).

VOLUME 94, NUMBER 2 Shi

Fig. 13. Zeuxo normani.

Material.—Velero IV _ sta. 23045 (33°53'02"N, 120°06’01”W), off Santa
Rosa Island, 55 m; 11 22:2 22 coll. Sieg, 2 2 2 in Senckenberg Museum,
Frankfurt, Germany; 7 2 2 in National Museum of Natural History, Wash-
ington: USNM 170655. Velero IV sta. 24358/BLM 636 (32°40'0’N,
119°9’49”W), Tanner Bank, southern California borderland, 120 m; 1 @
AHF, University of Southern California. Santa Cruz Island, Scorpio Har-
bour, California, 13 m; 1 2°; Willis G. Hewatt coll. (USNM 86280). All
specimens are syntypes.

Discussion.—This species closely resembles Zeuxo normani (Richard-
son), but can be distinguished by the relative proportions of the articles of
antenna |. The first article of Anatanais is twice the length of the second
(rarely these proportions may vary from 3:1 to 2.5:1 in Zeuxo). There is
also slight variation in some setae and plumose hairs, but they appear to be
of no taxonomic value. The present distribution is only at the designated
stations off Santa Cruz Island, Santa Rosa Island, and Tanner Bank within
the southern California borderland.

338 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 14. Zeuxo paranormani.

Zeuxo normani (Richardson, 1901)
Fig. 13

Synonymy, see Sieg (1980).

Discussion.—In order to determine correctly the species of Zeuxo it is
necessary to examine the coxa of pereopod | (Fig. 13), the carpus of per-
eopods 2-4, and the lacinia mobilis of the mandibles (Fig. 13). Zeuxo nor-
mani is Characterized by the triangular protuberance extending from the
coxa of pereopod 1, the 4 lateral and 2 medial spines borne distally on the
carpus of pereopod 2 and the slightly reduced lacinia mobilis of the right
mandible. In addition, the uropods (Fig. 13) of the adults consist of the basis
and an endopod of 5 articles (‘‘6-jointed’’). A detailed redescription of the
species is given, with figures, by Sieg (1980). The distribution of Z. normani
is difficult to determine due to confusion with Zeuxo paranormani. At pres-
ent it is reported from: Japan, without detailed locality (Shiino, 1951), and
Tanabe Bay (Bieri and Tokioka, 1968); British Columbia, Nanoase Bay,
False Narrows (Fee, 1927); Monterey Bay, California (type-locality, Rich-
ardson, 1905); Catalina Island, Anacapa Island, Santa Barbara Harbor,
Moss Beach (Miller, 1968), and La Jolla (USNM 53846, det. Sieg).

VOLUME 94, NUMBER 2 339

Fig. 15. Zeuxo coralensis.

Zeuxo paranormani Sieg, 1980
Fig. 14

A detailed description and discussion of this species, including figures,
may be found in Sieg (1980).

Discussion.—Zeuxo paranormani has been confused with Zeuxo nor-
mani. The adults of Z. paranormani may be distinguished by the presence
of uropods of 5 articles, while Z. normani have uropods of 6 articles (°*6-
jointed’’). The protuberance of the coxa of pereopod 1 is somewhat less
prominent in Z. paranormani than in Z. normani, and the carpus of per-
eopod 2 bears 2 medial and 4 lateral spines in Z. normani. The distribution
of the species is presently known from scattered localities along the Cali-
fornia coast: Humboldt Bay (USNM 66709), Santa Cruz Island (USNM
86281), La Jolla (USNM 53847), and Monterey Bay (USNM 30615), where
it is noted as Tanais normani Richardson, 1905 type-material.

Zeuxo coralensis Sieg, 1980
Fig. 15

Discussion.—This species has been mentioned as occurring along the Pa-
cific coasts of Japan and Panama Canal (Paitila, Sieg, 1980). It is included

VOLUME 94, NUMBER 2 34]

here because it possibly may occur along the California coast. For a dis-
cussion of the species see Sieg (1980).

Zeuxo maledivensis Sieg, 1980
Fig. 16

Discussion.—This species may also occur along the California coast, hav-
ing been noted from the Maldives, Japan, and Florida. It may be distin-
guished from the other species in the Tanaidae by the figured details (Fig.
16). For a detailed description and discussion see Sieg (1980).

Zeuxo seurati Nobili, 1906
Fig. 17

Discussion.—Zeuxo seurati 1s recognized by the reduction of the left and
right lacinia mobilis of the mandibles. It has been previously reported from
the Tuamotu Islands, Japan (Nobili, 1906) and the Hawaiian Islands as An-
atanais insularis (Miller, 1940).

Acknowledgments

The authors would like to express their appreciation to the Deutsche
Forschungsgemeinschaft and the Allan Hancock Foundation for their gen-
erous support of this work; to the United States Bureau of Land Manage-
ment for making specimens available; and to Dr. Richard C. Brusca, Dr.
Gilbert F. Jones, Dr. Charles G. Messing, and Dr. Thomas E. Bowman for
their critical reviews of the manuscript. This paper is Allan Hancock
Foundation contribution number 388.

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. 1962. The systematics and biology of bathyal and abyssal Isopoda Asellota.—Gala-

thea Reports 6:7—320.

(JS) Universitat Osnabriick, Abteilung Vechta, 2848 Vechta (Oldb.), Driv-
erstrasse 22, Postfach 1349, West Germany; (RNW) Allan Hancock Foun-
dation, University of Southern California, University Park, Los Angeles,
California 90007.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 344-351

A REDESCRIPTION OF ECHINASTER MODESTUS
PERRIER, 1881 (ASTEROIDEA: SPINULOSIDA)
FROM THE EASTERN GULF OF MEXICO
AND THE WEST INDIES

Richard L. Turner and David B. Campbell

Abstract.—The seastar Echinaster modestus Perrier, 1881, is redescribed
and a lectotype is designated. Echinaster spp. from the west coast of Florida
used in recent experimental studies are not E. modestus.

During a morphological study of seastars of the genus Echinaster from
the west coast of Florida, it was necessary for us to determine the identities
of two common sibling species that have been called E. spinulosus (Fer-
guson, 1966, 1967, 1968), E. echinophorus (Ferguson, 1970, 1974; Atwood,
1973a, 1973b, 1973c; Atwood and Simon, 1973), E. modestus (Turner, 1976;
Blake, 1978, 1980; Turner and Lawrence, 1979), and Echinaster (Ferguson,
1975a, 1975b, 1976, 1980). Based on the most recent monograph of seastars
of the region (Downey, 1973), both forms must be assigned to E. modestus
(Ferguson, 1975a, 1976; Turner, 1976; Turner and Lawrence, 1979); but a
recent note (Campbell and Turner, 1979) gave three distinct morphological
and meristic differences in the skeleton of the two forms. Examination of
type-specimens by us has revealed that the two forms and Downey’s (1973)
material do not belong to E. modestus. Perrier’s (1881, 1884) brief descrip-
tions of E. modestus might have contributed to the present confusion, and
Verrill’s (1915) description of Perrier’s specimens was equally brief. Wal-
enkamp’s (1979) detailed description seems to be based partly on material
that differs from Perrier’s type-specimens. Herein we redescribe E. modes-
tus from the type material and designate a lectotype.

Family Echinasteridae Verrill, 1867
Genus Echinaster Miller and Troschel, 1840
Echinaster modestus Perrier, 1881
Figs. 1-2

Echinaster modestus Perrier, 1881:7, 1884:206-207, pl. 3, fig. 7 (part).— |
Verrill, 1915:43-44 (part).—Clark, 1941:55.—Walenkamp, 1979:74-79
(part). |
Material.—MCZ 1080, Lectotype, | specimen (R = 34 mm, r = 8 mm),

Blake Sta. 45 (101 fm, 25°33’N, 84°21’W).—MCZ 1119, Paralectotype, 1

VOLUME 94, NUMBER 2 345

Fig. 1. Echinaster modestus, MCZ 1080, Lectotype. A, Aboral view; B, Adoral view.

346 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Echinaster modestus, planar views of ray ossicles from midline of aboral surface
(top) to the ambulacrum (bottom), near base of ray. Ossicles of primary skeleton unstippled;
ossicles of secondary skeleton (except spines) stippled. Rows of ossicles, top to bottom: car-
inals, accessory ossicles, adradials, dorsolaterals, superomarginals, intermarginals, inferomar-
ginals, actinolaterals, adambulacrals. Abbreviations: d, distal; p, proximal. A, MCZ 1080,
Lectotype, full spination of adambulacrals 12-15 and actinolaterals shown; B, MCZ 1119, Para-
lectotype, only furrow spines of adambulacrals 12—17 shown.

specimen (R = 22 mm, r = 6 mm), Blake Sta. 157 (120 fm), Montserrat.—
MCZ 4668, Paralectotype, 1 specimen (R = 26 mm, r = 8 mm), Blake Sta.
45 (101 fm, 25°33’N, 84°21’W).—MCZ 1081, 2 specimens (R = 9.5 mm, r
= 2.7 mm; R = 8.1 mm, r = 2.9 mm), Blake Sta. 272 (76 fm) and 292 (56
fm), Barbados.—MCZ 1118, 1 specimen (R = 5.7 mm, r = 2.0 mm), Blake
Sta. 276 (94 fm), Barbados.—MCZ 4001, 1 specimen (R = 56 mm, r = 11
mm), Atlantis Sta. 3303 (260 fm), Playa Baracoa, Cuba.

Diagnosis.—Rays 5, tapered, only slightly inflated at bases. Madreporite
circular, with radiating gyri; peripheral spinules present. Aboral skeleton
reticulate, with large papular areas. Adoral skeleton compact, with few pap-
ulae. Serial ossicles of rays: carinals, adradials, several rows of dorsolat-
erals, superomarginals, | short row of intermarginals, inferomarginals, 1-2

VOLUME 94, NUMBER 2 347

rows of actinolaterals, adambulacrals; all series except latter 2 with proximal
imbrication; glassy tubercles present. Spines small, blunt, conical; generally
1 per ossicle, except for superomarginals (1-2), inferomarginals (2—4), adam-
bulacrals (4-6), and mouth plates (6-7).

Description of type-material.—The following is based largely on MCZ
1080. Many details were confirmed in the two paralectotypes.

The rays taper uniformly to acute tips, with slight or no inflation near the
bases. The smaller type-specimens are more robust (R:r, 3.7 and 3.3) than
the larger (R:r, 4.3). The small, circular madreporite has a radiating system
of sulci and gyri and bears 6—13 spinules on its periphery. Anal spinules
number 8—14. The primary circlet has 1 (rarely 2) accessory ossicle between
adjacent primary radials and interradials, but there is none between the
madreporite and the primary radials of the bivium. Each primary radial
bears 1—2 (usually 1) small, blunt spines. Other ossicles of the primary circlet
and periproct bear 0-1 spine each.

The aboral skeleton is a loosely imbricating network. Papular areas are
wider than or equal in diameter to the width of surrounding ossicles. Papulae
are most abundant (3—7) between the carinals and adradials. The primary
series of ossicles on the aboral surface are the carinals and adradials, each
member of which bears a spine and overlaps the next proximal member
(proximal imbrication). Members of the carinal series are triangular and
alternate right and left of the midline; they are rarely separated by accessory
ossicles. Accessory ossicles sometimes join the carinal and adradial series,
which are otherwise directly joined. Members of the adradial series are
triangular or quadrangular and are not separated by accessory ossicles. A
few adradials have patches of glassy tubercles, but they appear eroded,
perhaps from storage. The major secondary elements of the aboral skeleton
are the dorsolaterals. Near the base of the ray, 3—S dorsolaterals form a
transverse series from each superomarginal to an adradial. The transverse
series of dorsolaterals are joined by 1-2 longitudinal series of more super-
ficial ossicles. A spine occurs on many members of the longitudinal series
and on some of the transverse members, forming 1-2 irregularly spinous
rows that extend */; the length of the ray.

The adoral skeleton is compactly imbricate, with few papulae. Supero-
marginals and inferomarginals are quadrangular ossicles with broadly round-
ed corners. They bear eroded patches of glassy tubercles and have proximal
imbrication. The 2 marginal series are separated at the base of the ray by
a few intermarginals, each of which has 0-1 spine. Intermarginals have
proximal imbrication. The superomarginals define the ambitus; they bear 1-
2 spines and are about as wide as they are long. The inferomarginals bear
2—4 spines and are wider than they are long. Each adambulacral bears a
transverse row of 4—6 spines: | (rarely 2) slender, webbed spine deep in the
ambulacral groove; | long, blunt spine at the margin of the groove (furrow

348 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

spine); a slightly shorter, more distal furrow spine; and 1-2 (rarely 3) short
subambulacral spines. Each mouth plate has 1 preoral spine, 3 furrow
spines, 0—1 (usually 1) suboral spine, and 1-2 spines within the groove and
hidden by the furrow spines. Actinolaterals form a distinct row between the
inferomarginals and adambulacrals for less than half the ray length and then
become hidden by the inferomarginals; each has 0-1 spine. A few additional
actinolaterals sometimes occur in a short row next to the inferomarginals,
but they are confined to the disc.

Description of other material.—Three other, small specimens from Blake
stations 272, 276, and 279 also are E. modestus. Even the smallest has
proximal imbrication, peripheral spinules (2) on the madreporite, 1-2 dor-
solaterals in transverse series, a few interbrachial intermarginals, and a row
of actinolaterals. The largest specimen is less than half the size of the small-
est type-specimen, but it agrees with the description above in every detail
except for the following: 3 peripheral spinules on madreporite; sulci and
gyri of madreporite poorly developed; number of anal spinules and nature
of primary circlet undetermined; few members of the longitudinal series of
dorsolaterals present; proportion of ray length occupied by dorsolaterals
undetermined; intermarginals not yet imbricate; superomarginals longer
than wide; inferomarginals as wide as long; mouth plates with only 2 furrow
spines. Most differences probably reflect ontogenetic variation. The third
specimen has many superficial characteristics of E. modestus, but the skin
was not removed for closer examination. As in the type-material, the smaller
specimens are progressively more robust (R:r, 3.5, 3.0, 2.9).

Clark’s (1941) large specimen from Atlantis Sta. 3303 seems to be E.
modestus with a few differences that might only reflect growth of the rays:
the carinal series and primary circlet have more accessory ossicles; the
intermarginals and actinolaterals extend % the length of the ray; calcareous
nodules lie free in the papular areas; rays are proportionately longer (R:r,
5x).

Type-locality.—Southeastern Gulf of Mexico, northwest of Dry Tortugas.

Discussion.—Perrier (1881) did not specifically designate the type-mate-
rial of Echinaster modestus, but he gave measurements for the larger spec-
imen from Blake Sta. 45 and published a photograph of it in 1884. He later
(1884, 1894) listed specimens from Gulf of Mexico, Montserrat, St. Vincent,
and Guadeloupe stations of the Blake but not the specimens from Barbados.
The material examined here represents all E. modestus held by the Museum
of Comparative Zoology, Harvard University. We have not seen the spec-
imens from St. Vincent and Guadeloupe. Because of the unsettled state of
systematics of Echinaster (Downey, 1973; Tortonese and Downey, 1977;
Tortonese and Madsen, 1979), we have designated the larger syntype from
Blake Sta. 45 as lectotype and the other syntypes (MCZ 1119 and MCZ
4668) as paralectotypes.

VOLUME 94, NUMBER 2 349

Our observations confirm Perrier’s (1881) brief diagnosis, but his descrip-
tion of spination around the ray did not reveal the kinds of underlying os-
sicles. The type-specimens differ from Downey’s (1973) description by the
presence of dorsolaterals, actinolaterals, proximal imbrication, and more
spines on the mouth plates; examination of some of her material confirmed
that it does not belong to E. modestus. Walenkamp (1979) examined some
type-specimens of FE. modestus as well as new material collected from the
Guyana Shelf. His description included the lack of glassy tubercles, 3 rows
of actinolaterals, a long row of intermarginals, and 4 mouth spines within
the furrow; these characteristics are not found in the type-material. He did
not describe the arrangement of dorsolaterals, but his description of ‘‘open
areas’ on the aboral surface probably reflects the arrangement of dorsolat-
erals and agrees with the type-material. Walenkamp (1979) assigned Down-
ey’s (1973) specimens of Verrillaster spinulosus to E. modestus. Our ex-
amination of her specimens, similar in size to the lectotype, revealed that
they have no glassy tubercles, too many intermarginals, and 2-3 rows of
actinolaterals. The identity of Downey’s specimens of V. spinulosus re-
mains unknown. Echinaster spp. on the west coast of Florida that have
been used for experimental studies cited above lack actinolaterals, have a
longer, less compact series of intermarginals, have fewer dorsolaterals and
oral furrow spines, and have distal imbrication (proximal ossicles Overlap
distal ossicles); they are not E. modestus.

Acknowledgments

We thank Dr. R. M. Woollacott, Ms. C. Felton, and Ms. S. Hunt, Mu-
seum of Comparative Zoology, Harvard University, and Dr. D. L. Pawson
and Ms. M. E. Downey, National Museum of Natural History, Smithsonian
Institution, for technical assistance and for the loan of material in their care.
Ms. A. M. Clark, British Museum (Natural History), gave helpful comments
on the manuscript. This study was supported in part by NSF grant SER 77-
06567.

Literature Cited

Atwood, D. G. 1973a. Larval development in the asteroid Echinaster echinophorus.—Biol.
Bull. 144:1-11.

- 1973b. Correlation of gamete shedding with presence of neurosecretory granules in
asteroids (Echinodermata).—Gen. Comp. Endocrinol. 20:347-350.

. 1973c. Histological comparison of asteroid neurosecretory granules before, during,
and after natural spawning.—Trans. Amer. Micros. Soc. 92:277—280.

, and J. L. Simon. 1973. Histological and histochemical analysis of neurosecretory
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| Biol. Soc. Wash. 91:234-241.

- 1980. On the affinities of three small sea-star families.—J. Nat. Hist. 14: 163-182.

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Campbell, D. B., and R. L. Turner. 1979. Comparative skeletal and ossicle morphology of ©
sibling species of Echinaster (Echinodermata: Asteroidea) from the west coast of Flor-
ida.—Amer. Zool. 19:1009.

Clark, H. L. 1941. 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.
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Downey, M. E. 1973. Starfishes from the Caribbean and the Gulf of Mexico.—Smithson.
Contrib. Zool. 126:1—158.

Ferguson, J. C. 1966. Mechanical responses of isolated starfish digestive glands to metabolic

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. 1970. An autoradiographic study of the translocation of amino acids by starfish.—

Biol. Bull. 138:14—25.

. 1974. Growth and reproduction of Echinaster echinophorus.—Fla. Sci. 37:57-60.

. 1975a. The role of free amino acids in nitrogen storage during the annual cycle of a

starfish.—Comp. Biochem. Physiol. 51A:341-350.

. 1975b. Fatty acid and carbohydrate storage in the annual reproductive cycle of Echi-

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. 1980. Fluxes of dissolved amino acids between sea water and Echinaster.—Comp.

Biochem. Physiol. 65A:291-295.

Perrier, E. 1881. Reports on the results of dredging under the supervision of Alexander Agas-

siz, in the Gulf of Mexico, 1877-78, by the United States Coast Survey Steamer

‘‘Blake,’’ Lieut.-Commander C. D. Sigsbee, U.S.N., commanding, and in the Caribbean

Sea, 1878-79, by the U.S.C.S.S. ‘‘Blake,’’ Commander J. R. Bartlett, U.S.N., com-

manding. 14. Description sommaire des Espéces nouvelles d’Astéries.—Bull. Mus.

Comp. Zool., Harvard 9:1-31.

. 1884. Mémoire sur les étoiles de mer recueillies dans la Mer des Antilles et le Golfe

du Mexique durant les expéditions de dragage faites sous la direction de M. Alexandre

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Turner, R. L. 1976. Sexual difference in latent period of spawning following injection of the.

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tebrates. Univ. S. Carolina Press, Columbia.

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those obtained by the Bahama Expedition from the University of Iowa in 1893.—Bull.
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VOLUME 94, NUMBER 2 351

Walenkamp, J. H.C. 1979. Asteroidea (Echinodermata) from the Guyana Shelf.—Zool. Verh.
170: 1-97.

Department of Biological Sciences, Florida Institute of Technology, Mel-
bourne, Florida 32901; (DBC) present address: Department of Zoology,
University of Rhode Island, Kingston, Rhode Island 02881.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 352-362

THE MORPHOLOGY OF TRIPOLYDORA SPINOSA
WOODWICK (POLYCHAETA: SPIONIDAE): AN
APPLICATION OF THE SCANNING ELECTRON
MICROSCOPE TO POLYCHAETE SYSTEMATICS

James A. Blake and Keith H. Woodwick

Abstract.—The morphology of Tripolydora spinosa Woodwick is reex-
amined using traditional light microscopy and the Scanning Electron Micro-
scope (SEM). In addition to its already-known unique features, T. spinosa
possesses modified spines on setiger 4, bristles on the major spines of setiger
5, unique ‘“‘ribbed’’ notosetae on setigers 7-10, and neuropodial inferior
sabre setae which accompany the hooded hooks. The setal arrangements,
as revealed by SEM, permit a consideration of possible setal homologies of
the modified spines of setigers 4 and 5. The position of Tripolydora among
those genera having a modified fifth setiger is considered, relative to other
genera of the subfamily Spioninae.

Tripolydora spinosa was first described from rocky intertidal habitats on
Enewetak Atoll in the Marshall Islands by Woodwick (1964). Kohn and
Lloyd (1964) subsequently reported the species from Easter Island where
it occurred in rocky tidal pools with sand and algae.

In the original description, Woodwick (1964) found 7. spinosa to possess
an unusual combination of characteristics including a partially modified fifth
setiger and branchiae on anterior setigers including the modified fifth. The
species also has a characteristic unique for a member of the Polydora-com-
plex, the presence of tridentate hooded hooks.

Because of this unusual combination of characteristics and the possible
significance of T. spinosa in the evolution of the Polydora-complex (Blake,
1979), the species was selected to be the subject of a more complete mor-
phological analysis. A detailed study of the morphology of T. spinosa using
both the light and Scanning Electron Microscope (SEM) was initiated. For
this work, it was possible to reexamine the Enewetak and Easter Island
specimens, all of which are deposited in the National Museum of Natural
History (USNM). In addition, new materials from Aitutaki in the Cook
Islands, collected by the first author (JAB), Johnston Atoll, provided by the
Naval Oceanographic Sorting Center (NOSC) in Hawaii, and from the Sey-
chelles (Indian Ocean), deposited in the USNM were also examined.

The species is redescribed. A discussion considers the homologies of setal
arrangements in forms having a modified fifth setiger and possible evolu-

) in right lateral view; C, Posterior end in dorsal view.

VOLUME 94, NUMBER 2

Bice. oe LE) Dy
6 LE.

aie, \
a ENN
8, wih

fn pion Nt 5 Goa
ee

Gee yl
oa
DISS. ae,

—

a unt pee 2 ae
SAWN CERRO

OT AVN 5 oto
Pian = aoe \\

We

le

Boa ° A A ( é
: 200 7 Nis —~ 2°

353

Fig. 1. Tripolydora spinosa (USNM 49530): A, Anterior end in dorsal view; B, Setigers 3-

onary significance of T. spinosa relative to its placement in the Polydora-
complex of the subfamily Spioninae.

Methods

Nomarski Differential Interference Contrast (Zeiss RA) and Phase Con-
rast (Zeiss GFL) optics were used for light microscopy. SEM studies were
made with a Zeiss Novascan 30 and JOEL JSM 35. SEM photographs were
oroduced from negatives of Polaroid Type 55 film. The specimens were all
rom previously fixed museum specimens which are ordinarily fixed in for-
nalin and preserved in 70% Ethanol. The specimens were dehydrated for
SEM observation in a critical point drier and sputter coated with gold-pal-
adium.

354 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Tripolydora spinosa (USNM 49530): A—C, Modified setae from setiger 4; D, Second
tier capillary notoseta from setiger 3; E-F, Major spines from setiger 5; G, Companion bi-
limbate capillary setae from setiger 5; H, Anterior tier capillary notoseta from setiger 7; I,
Posterior tier “‘ribbed’’ notoseta from setiger 7; J, Tridentate hooded hook; K, Fascicle of two
bidentate hooded hooks, two capillary neurosetae and an inferior sabre seta.

Systematics

Tripolydora spinosa Woodwick, 1964
Figs. 1—S

Tripolydora spinosa Woodwick, 1964:155-156, fig. 4(6—9).—Reish,
1968:222.—Kohn and Lloyd, 1973:700.

Material examined.—MARSHALL ISLANDS: Enewetak Atoll, Bogam-
bo, 2 July 1957, in beach rock, coll. D. J. Reish, holotype (USNM 32610).—
EASTER ISLAND: between Hanga Roa and Hanga Piko, 15 Feb. 1969, in

VOLUME 94, NUMBER 2 355)

Fig. 3. Tripolydora spinosa (USNM 49530): A, SEM of notopodia of setigers 4 and 5 in
right lateral location; B, SEM of same in posterior view of setiger 5. Scale: Fig. 3A = 20 um;
3B = 10 um.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Tripolydora spinosa (USNM 49530): A, SEM of modified notosetae (row I) and
capillaries (row II) from setiger 4 in posterior view; B, Major spines of setiger 5 in lateral
orientation. Scale: Fig. 4A = 10 wm; 4B = 10 um.

VOLUME 94, NUMBER 2 S)5)/

rock in tide pool, coll. A. Kohn and M. Lloyd, 30 specimens (USNM 49537);
same locality, in sand and algae, 125 specimens (USNM 49530).—COOK
ISLANDS: Aitutaki, Akiami motu, south side, 1 Aug. 1978, boring into
dead coral, coll. J. A. Blake, 3 specimens (JAB).—JOHNSTON ATOLL:
Sta. 3, transect 6, 25 April 1976, coll. R. Brock, 11 specimens (NOSC
77003J1).—_SEYCHELLES: Round Island, 19°45’S; 57°50’E, intertidal,
from sponge, Dec. 1975, coll. K. Buchanan, 3 specimens (USNM 599726).

Description.—This is a small species, measuring up to 3.5 mm long and
0.4 mm wide for about 37 setigerous segments. In alcohol the color is opaque
white to light tan with no body pigment. The prostomium is entire and
broadly rounded on the anterior margin (Fig. 1A). The caruncle is broad
and extends posteriorly over setigers 2 and 3 (Fig. 1A). There is no occipital
tentacle and no eyes. Nuchal cilia occur in long tracts on both sides of the
caruncle, merging posteriorly with the first dorsal sense organ on setiger 4.
The peristomium is well developed and broad, but not inflated.

Setiger 1 is greatly reduced and lacks a notopodium and notosetae. The
neuropodium is short, conical, and bears delicate capillary setae (Fig. 1A).
Setigers 2-10 have elliptically shaped notopodial lamellae and indistinct neu-
ropodial lamellae.

Setigers 2-10 bear notopodial fascicles of setae arranged in 2 rows (I and
II), each running in a general dorso-ventral orientation, and a third group
(III) of long superior capillaries located at a point near the uppermost setae
of the second row. The setae vary in length and structure. The setae of the
anterior row (I) are shortest, while those of the superior group (III) are the
longest. The setae of setigers 2 and 10 are the least modified in the sequence,
while those of setigers 4 and 5 are the most modified. There is a gradation
of setal structure in setigers 3 and 4 leading to setiger 5 and a corresponding
gradation in setigers 6—10 leading away from setiger 5.

The notosetae of setigers 2—3 and 6 are unilimbate capillaries (Fig. 2D) in
which the setae of row I are shorter and have a broader sheath. In setigers
3 and 6 these setae are pennoned, with a long tapered tip. The enlarged
portion of these setae is seen by SEM to be composed of free surface fibrils
which are longest on one side of the seta. Varied lengths of these fibrils
produce a tapered tip, which superficially resembles the acuminate setae of
sabellids.

The notosetae of setiger 4 are arranged as on setigers 2 and 3 except that
row I is modified and bears 3-4 setae having inflated sheaths, through which
the tip of the shaft often protrudes (Figs. 2A—C; 3A—B; 4A). When observed
with the SEM, the inflated sheath is seen to be a crest of bristles (Fig. 4A).

Setiger 5 appears only slightly modified superficially (Fig. 1A—B). The
setae include a diagonal row of 3-4 major spines corresponding to row I and
positioned anteroventral to 2 groups of 2—3 bilimbate companion capillaries
(Fig. 2G) derived from row II, and a fascicle of dorsal capillaries (Fig. 3A—

358 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Tripolydora spinosa (USNM 49530): A, SEM of “‘ribbed’’ row II capillary notosetae
from setiger 7 in posterior view; B, Detail of same; C, SEM of tridentate hooded hooks in
lateral view; D, Detail of individual hook in frontal orientation. Scale: Fig. SA = 4 wm; 5B
= 5 yum; 5D = | pm.

VOLUME 94, NUMBER 2 359

B) corresponding to row III. The major spines have a terminal tooth flanked
on one side by 2 short knobs; the central depression between these prom-
inences bears fine bristles (Fig. 2E; 3B; 4B). Worn spines have reduced
knobs (Fig. 2F).

Setigers 7-10 have short unilimbate capillary notosetae in row I (Fig. 2H)
similar to those of setigers 2—3 and 6 and long unilimbate capillaries with
unusual riblike sculptured sheaths (Fig. 2I) in row II. The SEM reveals
these ribs to consist of thickened whorls of minute bristle endings (Fig. 5A—
B). The notosetae become longer and more slender in subsequent setigers,
with those of the far posterior setigers being arranged as dense bundles of
long needles which project posteriorly over the dorsum (Fig. 1C).

The neurosetae of setigers 2—8 consist of 4-5 unilimbate capillaries. Tri-
dentate hooded hooks begin on setiger 9 and number 2—4 per neuropodium
throughout the body. Initially, the hooks are accompanied by 4—5 unilimbate
capillaries, but these are gradually replaced by 2-3 simple capillaries and a
single large inferior sabre seta (Fig. 2K) bearing granulations near the distal
end. The hooks have a large and a small apical tooth surmounting the main
fang, to which the large apical tooth is closely applied (Fig. 2J). The small
apical tooth is often worn and sometimes lacking, especially in posterior
setigers (Fig. 2K). With the SEM, the hood is seen to have a fine micro-
structure, consisting of numerous minute bristle endings (Fig. 5C—D). In
some cases, the hood is inflated (Fig. 5C). The hood opening is expanded
near the tip of the main fang and flares somewhat laterally (Fig. SD).

Branchiae occur from setiger 2, and continue without interruption to near
the posterior end. The branchiae on setigers 2—5 are shorter than those of

‘setiger 6 and subsequent setigers (Fig. 1A). Dorsal sense organs begin on
setiger 4 and extend across the segments between the gills (Fig. 1A). The
dorsal sense organs are difficult to see with light microscopy, but prominent
with SEM. The pygidium is formed by 4 small lobes, the dorsal pair smaller
(Fig. 1C).

Distribution.—Pacific Ocean: Marshall Islands, Johnston Atoll, Cook Is-
lands, Easter Island.—Indian Ocean: Seychelles.

| Discussion

The SEM reveals a significant number of details about setal structure and
ciliary arrangement of Tripolydora spinosa that are not apparent with the
light microscope. The SEM also demonstrates that it is possible to interpret
setal homologies by considering setal arrangements over successive seg-
‘Ments.
| The arrangement of modified spines and capillaries on setigers 4 and 5 is
clearly revealed in Fig. 3A. The modified spines of setiger 4 appear to be

360 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

homologous to the shorter and thicker unilimbate capillaries occurring in
row I of setigers 2-3 and 6-10. The same homology holds for the major
spines of setiger 5. Here the 3—4 spines are aligned in a more anteroventral
position. The long, thin capillaries of row II are also shifted on setigers 4
and 5. On setiger 4, the 2-3 ventralmost and shortest capillaries lie just
posterior to the modified setae. This same relationship occurs on setiger 5,
with the ventralmost capillaries becoming the companion setae.

The modified notosetae of setiger 4 are unusual. In some respects they
resemble the major spines of setiger 5 found among some species of Poly-
dora and Boccardia where the curved shaft is surmounted by a crest of
bristles. In T. spinosa this crest is transparent when viewed with the light
microscope, but is a solid feature in Polydora caulleryi Mesnil (see Blake,
1971). It is probable that the modified notosetae on setiger 4 of 7. spinosa
are transitional in degree of modification between unilimbate capillaries and
the heavy crested spines of P. caulleryi.

The major spines of setiger 5 on T. spinosa are revealed by SEM to have
a terminal tooth and 2 shorter knobs between which are located fine bristles.
The terminal structure of this seta is very difficult to observe with light
microscopy and the bristles are rarely seen. This emphasizes the difficulties
of making accurate portrayals of setal structure using only the light micro-
scope. The bristles had not been previously reported on the major spines
of T. spinosa.

The ribbed nature of the capillary notosetae of setigers 7-10 of T. spinosa
has not been previously described. With light microscopy these setae have
a unilimbate appearance in which the ribs appear to be surface sculptures
of the sheath. With SEM, it can be seen that each “‘rib”’ is part of a whorl
of thickened fibril endings which completely cover the ends of these setae.

The fine details of the hoods of the hooded hooks are different in T.
spinosa than we have observed using SEM in related species of Polydora
and Boccardia. These differences manifest themselves in the degree of bris-
tling apparent upon the hood, position and shape of the hood opening, and
the number of teeth which project through the hood opening. These differ-
ences suggest that microarchitecture of the hooded hooks may prove to be
useful in systematic studies of spionid polychaetes. Readers should be
aware, however, that hood form will differ depending upon the drying tech-
nique employed during specimen preparation. If the specimens are air dried
prior to coating and observation, the hoods tend to collapse, tear and more
closely adhere to the shaft and teeth. The bristles also tend to be distorted
with air drying. With critical point drying the hoods remain inflated, are
seldom torn, and the bristles remain undistorted.

Among the genera of the Polydora-complex, Tripolydora spinosa is
unique in many respects. It is the only species to have tridentate hooded

VOLUME 94, NUMBER 2 361

hooks, prominent granulated inferior sabre setae, and unique “‘ribbed’’ no-
tosetae on setigers 7-10. Moreover, with the exception of Boccardiella lim-
nicola (Blake and Woodwick, 1976), it is the only species with gills from
setiger 2 continuing uniformly to near the end of the body. The species has
modified notosetae on setiger 4 in addition to the major spines on setiger
5. Modifications to notosetae on setiger 4 are known for some species of
Pseudopolydora (see Woodwick, 1964), but are not known for other genera
of the Polydora-complex. Tripolydora spinosa also resembles Pseudopo-
lydora in the lack of major parapodial alteration to setiger 5 and in the
reduced angle between the main fang and apical tooth (teeth) of the hooded
hooks. The modified notosetae of setiger 4, the “‘ribbed’’ notosetae of se-
tigers 7-10 and the inferior sabre setae were not reported by Woodwick
(1964) because some of these setae were sheared off the holotype.

Blake (1979) proposed that Tripolydora exhibits a close relationship to
the non-polydorid genus Microspio. This conclusion was originally based
on similarities in branchial distribution, hooded hook structure, and the
presence of inferior sabre setae in both genera. Additional support for this
conclusion is the recent discovery that Microspio granulata Blake and Ku-
denov from Australia, the Hawaiian Islands, and Ifaluk Atoll in the Caroline
Islands, and an undescribed species of Microspio from Ecuador have en-
larged capillary notosetae on rows I and II over setigers 4—6 (Blake, un-
published). In addition, the setae of these 2 rows come together ventrally,
close and form a loop. This arrangement is reminiscent of some species of
Polydora where the modified spines of setiger 5 are positioned into a J-
shape (see Blake, 1971). This trend in Microspio towards modification of
setae in the corresponding modified region of polydorids occurs in no other
member of the Spionidae. Blake (1979) emphasized that larval characters
are also useful in interpreting relationships of the Polydora-complex; how-
ever, the larval morphology of Tripolydora spinosa has not been described.

From the information available, it would appear that Microspio is an
appropriate genus to consider as having given rise to the Polydora-complex.
In order to pursue this line of research SEM studies are being undertaken
on selected species of Microspio, Pseudopolydora, and other genera of the
Polydora-complex. Following completion of those studies it should be pos-
sible to describe more fully the interrelationships of the genera of the
subfamily Spioninae.

Acknowledgments

We are grateful to Linda Ward, University of Hawaii, and Marian H.
Pettibone, Smithsonian Institution for providing the specimens upon which
this study is based. Nancy J. Maciolek read and commented upon the manu-
script.

362 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Literature Cited

Blake, J. A. 1971. Revision of the genus Polydora from the east coast of North America

(Polychaeta: Spionidae).—Smithsonian Contrib. Zool. 75:1-32.

. 1979. Evolutionary relationships of spionid polychaetes.—Amer. Zool. 19:887.

, and K. H. Woodwick. 1976. A new species of Boccardia (Polychaeta: Spionidae)

from two freshwater lakes in southeastern Australia.—Rec. Aust. Mus. 30:123-128.

Kohn, A. J., and M. C. Lloyd. 1973. Marine polychaete annelids of Easter Island.—Int. Rev.
ges. Hydrobiol. 58:691-712.

Reish, D. J. 1968. The polychaetous annelids of the Marshall Islands.—Pacific Sci. 22:208-—
231.

Woodwick, K. H. 1964. Polydora and related genera (Annelida, Polychaeta) from Eniwetok,
Majuro and Bikini Atolls, Marshall Islands.—Pacific Sci. 18:146—-159.

(JAB) Battelle New England Marine Research Laboratory, P.O. Drawer
AH, Duxbury, Massachusetts 02332; (KHW) California State University,
Fresno, Biology Department, Fresno. California 93740.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 363-373

SYSTEMATIC NOTES ON CERTAIN OSCINES FROM
PANAMA AND ADJACENT AREAS
(AVES: PASSERIFORMES)

Storrs L. Olson

Abstract.—Vireo gilvus disjunctus Zimmer, 1941, is synonymized with
Vireo gilvus dissors Zimmer, 1941, the range of which is extended to include
both slopes of the northern end of the Western Andes of Colombia and
Darién, Panama. The range of Chlorospingus ophthalmicus novicius Bangs,
1902, is restricted to the Boquete region and the east slope of Volcan de
Chiriqui, Panama, the remainder of western Panama and Costa Rica being
occupied by C. o. regionalis Bangs, 1906. Chlorospingus punctulatus Scla-
ter and Salvin, 1869, should be regarded as a subspecies of C. ophthalmicus
(Du Bus, 1847). Chrysothlypis chrysomelas titanota n. subsp. is named from
Costa Rica, the nominate form being restricted to western Panama. Icterus
chrysater hondae Chapman, 1914, is considered to be known from the ho-
lotype and paratype only, and all other specimens from Colombia and Pan-
ama are referred to J. c. giraudii Cassin, 1848. Icterus mesomelas salvini
Cassin, 1867, extends only to the Almirante region of the Caribbean slope
of western Panama, all other specimens from Panama being referable to /.
m. carrikeri Todd, 1917. Zonotrichia capensis orestera Wetmore, 1951, is
synonymized with Z. c. costaricensis Allen, 1891.

Vireonidae
Vireo gilvus

The collection of a specimen of Vireo gilvus from Darién, Panama, that
is quite distinct from the population of western Panama and Costa Rica, V.
g. chiriquensis (Bangs, 1903), necessitates a review of the subspecies known
from Colombia. The last revision of these forms is that of Zimmer (1941).
Material in the USNM collected by Carriker subsequent to Zimmer’s revi-
sion requires changes to be made in the current nomenclature and published
distributions of certain subspecies.

Two subspecies are recognized in eastern Colombia—V. g. mirandae
Hartert, 1917, of the Santa Marta region and northern Venezuela, and V. g.
leucophrys (Lafresnaye, 1844) in the Eastern Andes of Colombia, reportedly
extending as far south as the eastern part of central Peru (Blake, 1968). As
noted by Hellmayr (1935) and Meyer de Schauensee (1951), mirandae is
poorly marked, but in series the 23 USNM specimens from Santa Marta,

364 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Sierra Perija, and Sierra Negra, Colombia are seen to have paler underparts
than /eucophrys, although not all individuals would be separable.

The populations of the middle portions of the Central and Western Andes
were recognized by Zimmer (1941) as a new race, V. g. dissors (type lo-
cality, Cerro Munchique, Cauca). This can be distinguished from leucophrys
by the grayer, as opposed to warm brown, crown and the greener, less
brown, dorsum. Although Meyer de Schauensee (1951) referred specimens
from Huila to leucophrys, Carriker specimens from Huila (3 from Belén, 45
km SW of La Plata; 2 from La Candela, 10: miles SW of San Augustin) show
a decided approach to dissors and can probably be regarded as intergrades.
An older specimen from La Candela, presumably through foxing, appears
more typical of leucophrys than those taken 40 years later.

On the basis of six specimens from the eastern slope of the northern end
of the Central Andes, Zimmer (1941) described an additional race, V. g.
disjunctus (type locality, Santa Elena, Antioquia). USNM specimens from
the Central Andes in Antioquia include 4 from Hacienda Zulaiba, 17 miles
NE of Santa Rosa de Osos, and 2 from La Bodega, S side of Rio Negrito,
on the road from Sonson to Narino. Additional specimens from Antioquia
definitely extend the range of the species to both slopes of the northern end
of the Western Andes, whereas previously there had evidently been some
doubt that the species occurred there (Blake, 1968). USNM specimens from
the Western Andes include 7 from Hacienda Potreros, 15 miles SW of Fron-
tino; 3 from Hacienda La Illusion, Rio Urrao, base of Paramo de Frontino,
and | from Urrao.

Although Zimmer (1941:18) took some pains to distinguish disjunctus
from the races in Ecuador and Peru, the only supposed difference from
dissors, which is the nearest race geographically, was given as ‘darker
upper parts.’ The characters he cites as differentiating disjunctus from
leucophrys are exactly the same as those given for separating dissors from
leucophrys. In the series I examined, which included topotypes of both
dissors and disjunctus, there was only a tendency for darker coloration in
the dorsum of more northern birds, but individual and seasonal variation
made it impossible to separate specimens into two groups. Therefore, one
of Zimmer’s names, which were proposed simultaneously, should be sup-
pressed. As first revisor, I select the name dissors to represent the subspe-
cies of Vireo gilvus inhabiting the entire extent of the Western and Central
Andes of Colombia, with disjunctus falling into synonymy.

A considerable extension of range for the species is provided by a single
female taken 4 June 1963 by Pedro Galindo, 4 miles (6.4 km) W of the
summit of Cerro Mali, Darién, Panama, at an elevation of 4800 feet (1463
m). This specimen (USNM 486488) has the pale underparts typical of Co-
lombian birds and cannot, therefore, be referred to chiriquensis of western
Panama. The coloration of the upperparts is unlike that of leucophrys and

VOLUME 94, NUMBER 2 365

is most like that of dissors, to which I tentatively refer the specimen, al-
though its preservation and state of wear do not permit a positive deter-
mination.

Both Hellmayr (1935) and Zimmer (1941) discuss a specimen of V. gilvus
from Ricaurte, Narino, southwestern Colombia, that Hellmayr referred to
v. g. josephae Sclater, 1859, of Ecuador. In the USNM collections is an
additional specimen, also from Ricaurte, taken by Carriker in 1950. I could
detect no difference between this specimen and individuals of dissors,
whereas it differs from josephae in the lighter crown and greener, less
brownish, dorsum. For the present, the distribution of V. g. josephae prob-
ably should not be considered to include Colombia.

Thraupidae
Chlorospingus ophthalmicus

There are two races of this highly polymorphic species recognized in
Costa Rica and Panama—C. o. novicius Bangs, 1902, and C. o. regionalis
Bangs, 1906. Examination of extensive material from Panama shows that
these two taxa are indeed valid, but their distribution and characters are not
properly delimited at present. Zimmer (1947:3), for example, considered
that “‘the separation of novicius and regionalis is not sharp and it is difficult
to say where the line should be drawn.’’ This idea results in part from
ambiguous labeling of specimens but also from the fact that novicius, which
is actually quite distinctive, has a peculiarly restricted range.

Bangs (1902) described Chlorospingus novicius from a series of specimens
collected by W. W. Brown in Chiriqui, western Panama. Most of these were
labeled as being from Boquete, but a few were labelled ‘“‘Mt. Chiriqui’’ or
‘‘Caribbean Slope,’’ meaning the Caribbean slope of Mt. Chiriqui (= Volcan
de Chiriqui or Volcan Baru). The holotype is from ‘‘Mt. Chiriqui’’ at 7500
feet (2286 m). It is important to note that Brown worked out of Boquete and
would therefore presumably have taken this, and other specimens so la-
beled, on the eastern slope of Volcan de Chiriqui. Bangs (1902) referred to
Boquete as being “‘south’”’ of Volcan de Chiriqui and considered most of
Brown’s collecting to have been done on the south slope, apparently to
contrast it with the “‘northern’’ or Caribbean slope, whereas in fact, Boquete
is more nearly east of Volcan de Chiriqui. Brown would have collected on
the eastern as opposed to the western face of the volcano, which latter has
been accessible only in more recent times. Because the birds from the west-
ern slope are referable to regionalis, the type locality of novicius should be
restricted to the eastern slope of Volcan de Chiriqui.

The species Chlorospingus ophthalmicus was poorly known when Bangs
(1902) described novicius, and he made his comparisons chiefly with the
subspecies from Bolivia now known as bolivianus Hellmayr, 1921, noting

366 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

particularly that the bill in novicius was larger. Bangs (1906) later described
the birds of Costa Rica as a new form, regionalis (type locality, Cariblanco
de Sarapiqui), distinguished by differences in color. Hellmayr (1936:402), in
contrasting novicius with regionalis, regarded it as having a *“much larger,
thicker as well as longer bill.’’ This is erroneous and was obviously adapted
from Bangs’ (1902) original comparison of novicius with the small-billed
race bolivianus. Hellmayr further considered that two specimens from
Urujas de Térraba, Costa Rica, were “‘identical with a topotypical series’’
of novicius, so that thenceforth (e.g. Storer, 1970), the range of novicius
has been given as including western Panama and southwestern Costa Rica,
with regionalis occupying the remainder of Costa Rica and Nicaragua.

In the process of arranging the USNM series of C. ophthalmicus from
Panama by locality to check geographic variation, I found that birds from
Boquete were markedly distinct from those taken elsewhere in Panama—so
much so that I was easily able to pick out the remainder of the Boquete
specimens before reading the labels. I could not distinguish birds from west
of Volcan de Chiriqui from regionalis of Costa Rica. Much has been obfus-
cated by the apparent tendency of collectors to label specimens taken any-
where within sight of Volcan de Chiriqui as “‘El Volcan.’’ Wetmore, for
example, applied this term to such diverse localities as Cerro Picacho (7 km
N of the peak), Silla de Cerro Pando (17 km WNW of the peak), and Lagunas
(16 km WSW of the peak). Fortunately, he also supplied the more detailed
locality information as well, which other collectors usually have not done.
Many specimen labels of the past few decades apparently refer to the town
of El Volcan (El Hato del Volcan or El Hato), west of the volcano, rather
than to the peak itself.

From the available evidence, it appears that C. 0. novicius has a much
more restricted distribution than hitherto suspected, being known with cer-
tainty only from the Boquete area and extending from there to the Caribbean
slope of Bocas del Toro. In addition to the USNM series of specimens from
Boquete, I have examined the holotype of C. 0. novicius and what remains
at the MCZ of the original type series (Mt. Chiriqui, 3; Caribbean slope, 2;
Boquete, 11), as well as the specimens recorded by Blake (1958) from the
Boquete area, including the Caribbean slope of Bocas del Toro. All of these
specimens are referable to novicius. Birds from the western slope of Volcan
de Chiriqui through Costa Rica to Nicaragua are referable to regionalis.

C. o. novicius differs from C. o. regionalis as follows: crown and sides
of head darker, decidedly brownish, not gray; dorsum more olivaceous, less
yellow-green; pectoral band more intensely orangish, less yellow; light por-
tions of throat more restricted, more heavily speckled, usually suffused with
buff or even with the orangish of the breast, not whitish as in regionalis.

In these characters, as Zimmer (1947) noted, novicius shows a decided
approach to the birds of Veraguas and Coclé, now known as Chlorospingus

VOLUME 94, NUMBER 2 367

punctulatus Sclater and Salvin, 1869 (see Storer, 1970), in which the crown
and sides of head are very dark brown, the throat is even more intensely
speckled, and the pectoral band is deeper orange. Both Zimmer (1947) and
Eisenmann (1955) rightly considered punctulatus to be but a subspecies of
C. ophthalmicus, but Storer (1970:259, footnote) quoted Wetmore (in litt.)
as believing it ‘“‘closer to [Chlorospingus | pileatus but would maintain it as
a full species until more is known about it in life.”’ C. punctulatus has the
white spot behind the eye characteristic of most subspecies of C. ophthal-
micus and shows no approach whatever to C. pileatus Salvin, 1864, which
has a broad white stripe extending from above the eye to the posterior
margin of the crown. In its coloration, punctulatus differs only slightly from
other dark-capped subspecies of C. ophthalmicus, such as venezuelanus
Berlepsch, 1893, and clearly should be regarded as a subspecies of C.
ophthalmicus. The fact that no specimens of C. ophthalmicus are known
from the area between Boquete and the range of punctulatus in Veraguas,
has no doubt contributed to the misunderstanding of the relationships of
punctulatus.

There is some evidence of intergradation between novicius and region-
alis. Three birds collected at Cerro Punta in 1932 show some buffiness in
the throat, a somewhat more orangish pectoral band, and some brown in
the crown, possibly due to intergradation with novicius. On the other hand,
two birds from the same locality taken in 1955, and another taken in 1965,
are clearly referable to regionalis.

USNM Panamanian specimens examined.—C. o. novicius: CHIRIQUI:
““Mt. Chiriqui’’ (topotypical paratype) 1; Boquete (paratypes) 4; Boquete,
Palo Alto, 1; Boquete, Rio Caldera, 5; Boquete, Quiel, 18.

C. o. regionalis: CHIRIQUI: ‘‘Volcan,’’ 2; El Volcan, Tisingal, 4; El
Volcan, Silla de Cerro Pando, 5; El Volcan, Quebrada Zumbona, 2; El
Volcan, Laguna Grande, 4; El Volcan, Lagunas, 2; El Volcan, Cerro Pica-
cho, 1.

Chrysothlypis chrysomelas

Currently there are two subspecies recognized for this distinctive bird.
The nominate form C. c. chrysomelas (Sclater and Salvin, 1869; type lo-
cality, Cordillera del Chucu, Veraguas, Panama) is stated to range through
Costa Rica and western Panama, and C. c. ocularis Nelson (1912; type
locality, Cana, Darién, Panama) has heretofore been known only from Da-
rién, eastern Panama. Nelson (1912) distinguished ocularis from chryso-
melas partly by the black loral spot of adult males, but mainly by what he
supposed to be differences in the females. He was misled by comparing
females of ocularis, which are entirely yellow underneath, with those from
Costa Rica, which he regarded as “‘typical’’ and which are whitish below.
Nelson had no specimens of true chrysomelas from western Panama, how-

368 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ever; these birds are entirely yellow below, as in ocularis. This is clearly
shown in Sclater and Salvin’s (1869) plate accompanying the original de-
scription of chrysomelas. The Costa Rican birds thus represent an unde-
scribed subspecies, as was mentioned in a footnote by Griscom (1935:377).

Chrysothlypis chrysomelas titanota, new subspecies

Holotype.—Female, USNM 209197, Bonilla, northern Cartago, Costa
Rica. Collected 10 April 1908 by Francisco Basulto, original number 743.

Characters.—Adult males like chrysomelas; females with throat and belly
suffused with white, leaving only a band across the breast yellowish, where-
as females of chrysomelas are entirely bright yellow below; flanks grayish,
not yellow-green as in chrysomelas.

Range.—Caribbean slope of Costa Rica. The great majority of specimens
come from a limited area in central Costa Rica, but Slud (1964:366) records
the species from “‘just south of Lake Arenal to the northern and eastern
sides of the Cordillera Central and the approaches to the Talamanca Cor-
dillera.. 2...

Etymology.—Greek, titanotos, whitened, from titanos, chalk or gypsum.

Specimens examined.—COSTA RICA. HEREDIA: Cariblanco de Sara-
piqui (adult ¢: 1, MCZ; 9: 1, MCZ; 1, AMNH). SAN JOSE: Carrillo (adult
6: 1, FM; 1, ANSP; 11, MCZ; immature ¢: 1, AMNH; 1, MCZ; 1, FM;
@: 2, MCZ; 4, AMNH); La Hondura (adult ¢: 1, USNM). CARTAGO:
Bonilla (adult ¢: 7, USNM; 2: 4, USNM; 1, AMNH).

Remarks.—The three immature males listed above are more yellow below
than any of the females. K. C. Parkes (in litt.) reports that of about 30
Specimens in the Carnegie Museum of Natural History (all from Carillo),
there are four non-juveniles sexed as females, two of which are whitish, as
typical for titanota, one of which is all yellow below, and one of which is
intermediate. One bird sexed as male that apparently is also a juvenile, is
whitish below, as is an unsexed juvenile. Thus there seems to be some
infrequent variation in the color of the underparts, but whether this can be
correlated with age or sex cannot be determined with the material at hand.
Regardless, virtually all Costa Rican females are instantly separable from
all Panamanian females.

Chrysothlypis chrysomelas chrysomelas (Sclater and Salvin)

Tachyphonus chrysomelas Sclater and Salvin 1869, Proc. Zool. Soc. Lon-
don 1869:440. Cordillera del Chucu, Veraguas, Panama.

Characters.—Females like ocularis, separable from titanota by the en-
tirely yellow underparts. Males like titanota, separable from ocularis by
the lack of a black loral spot.

VOLUME 94, NUMBER 2 369

Range.—Known with certainty only from Veraguas, Coclé, and western
Panama Province, Panama.

Specimens examined.—PANAMA. COCLE: Head of Rio Guabal (adult
6: 2, USNM; immature ¢: 2, USNM; 2, 2 USNM). PANAMA PROV-
INCE: Cerro Campana (adult 6; 1, USNM; 1, AMNH; 2: 1, USNM).

Remarks.—The identity of 8 specimens (MCZ) from the Caribbean slope
of Bocas del Toro, on the Boquete trail above Almirante (Kennard and
Peters, 1928; Peters, 1931), remains in doubt. Five are adult males, two are
immature males, and one is unsexed, but not an adult male. The last three
birds are entirely yellow below, as in chrysomelas, but in the absence of
any certain female specimens, the possibility remains that these birds may
be referable to titanota.

Chrysothlypis chrysomelas ocularis Nelson

Chrysothlypis chrysomelas ocularis Nelson 1912, Smithsonian Misc. Coll.
60:19. Cana, eastern Panama.

Characters.—Exactly like chrysomelas except adult males with part of
loral area black instead of yellow.

Range.—Eastern Panama in Darien and eastern Panama Province.

Specimens examined.—PANAMA. PANAMA PROVINCE: Cerro Azul
(adult ¢d: 1, AMNH). DARIEN: Cana (adult ¢: 2, USNM; immature ¢c: 1,
USNM; @: 2, USNM); Tacarcuna Village (adult ¢: 1, USNM; 2°: 2,
USNM); Cerro Tacaracuna (6: 2, AMNH; 2: 4, AMNH); Cerro Sapo
(3d: 5, ANSP; 2: 5, ANSP).

Remarks.—The supposedly greater width of the black orbital ring de-
scribed by Nelson (1912) is too much affected by the preparation of the
specimen to be of use. The specimen from Cerro Azul is the only one known
outside of Darién. It has the black loral spot somewhat reduced, but no
more so than certain individuals from Cerro Sapo.

Icteridae
Icterus chrysater

In the earlier literature, the southern populations of J. chrysater from
Panama, Colombia, and Venezuela, were considered to represent a single
subspecies, J. c. giraudii Cassin, 1898 (type locality **“Bogota’’). On the
basis of two (not three, contra Meyer de Schauensee, 1951) male specimens
from Honda, Tolima, Colombia, Chapman (1914) described a new species,
Icterus hondae, differentiated from J. chrysater by much deeper orange
coloration and supposedly smaller size. Meyer de Schauensee (1951), while
allowing that these specimens were more orange than in giraudii, considered
hondae to be a subspecies of chrysater. He assigned all birds from Panama,

370 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

the lowlands of western Colombia, and the Magdalena Valley to hondae on
the basis of supposedly smaller size, reserving the name giraudii for the
birds of the upper tropical and lower temperate zones of the remainder of
Colombia and part of Venezuela. Blake (1968) followed this treatment.

According to Meyer de Schauensee (1951), the wing is 90-92 mm in the
types of hondae, 89-94 mm in birds from the Pacific Coast of Colombia,
and 86—94 mm in specimens from Panama. He considered the wing length
of giraudii to range from 95.5 to 111 mm. Both the holotype and paratype
of hondae, however, retain the primaries of the previous subadult plumage,
which results in a shorter wing measurement. The wings in 32 males from
Panama in the USNM range from 89 to 103 mm, and average 96 mm. Nine
males from Magdalena and Norte de Santander, within the restricted range
of giraudii, had wing lengths from 96.3 to 101.6 mm, averaging 98.4 mm.
Overlap is thus far too great to permit separation of these birds, and there
are no size differences within the remainder of the USNM series from Co-
lombia and Panama that will permit the recognition of more than one sub-
species in this area. Alexander Wetmore’s unpublished notes and measure-
ments of this series indicate that he considered all these specimens to be
referable to J. c. giraudii, and I concur.

Although there is considerable variation in the intensity of coloration of
specimens of giraudii, none is as deep orange as the type and paratype of
I. hondae, which I examined at AMNH. Furthermore, the bills in these
specimens are proportionately longer, more slender, and pointed than in
giraudii; the mandibles in both are light horn color at the base, not bluish
or blackish as in giraudii; and both have less black on the forehead and
over the eye than giraudii. These characters were not noted in the original
description or subsequently. For the present I regard J. hondae as a prob-
lematical subspecies of J. chrysater that is known so far only from the two
original specimens. It is possibly confined to the upper Magdalena River
Valley.

Icterus mesomelas

Four subspecies are recognized in J. mesomelas, of which the northern-
most (mesomelas [Wagler, 1829]) and southernmost (taczanowskii Ridg-
way, 1901) are easily separated from the other two by the yellowish-white
margins of their inner secondaries. Birds from eastern Panama and the Ca-
ribbean slope of western Panama through Nicaragua are presently (Blake,
1968) referred to J. m. salvini Cassin, 1867 (type locality, Costa Rica),
whereas those from Colombia and western Venezuela are referred to J. m.
carrikeri Todd, 1917 (type locality, Fundacion, Santa Marta, Magdalena,
Colombia). Todd (1917) merely compared carrikeri with taczanowskii of
Ecuador and Peru and did not say how it differed from salvini. Hellmayr

- VOLUME 94, NUMBER 2 371

(1937) diagnosed carrikeri as differing from salvini in having the black gular
area more restricted, the bill smaller, and the general coloration slightly
more orange.

In analyzing the specimens in the USNM, it quickly became evident that
there were inaccuracies in the current concept of variation in this species,
as the majority of Panamanian birds proved inseparable from those of Co-
lombia. This at first suggested that the race carrikeri was invalid. Although
poorly marked, carrikeri can be salvaged by redefining its range to include
all of Panama east of the Canal Zone, extending west to Cerro Campana,
western Panama Province on the Pacific slope, and at least to El Uracillo,
Coclé, on the Caribbean slope. The only specimens from Panama that are
referable to salvini are from the Almirante area of Bocas del Toro (Almi-
rante 2, USNM; 5, MCZ; Western River 2, MCZ; Fruitdale 1, MCZ; Chan-
guinola River 2, MCZ). There is thus a considerable hiatus between salvini
and carrikeri along the Caribbean coast of Panama. This is in large measure
an artifact of collecting, but the pattern of distribution of these two subspe-
cies, as redrawn here, is one commonly observed in other polytypic species
of Panamanian birds.

Icterus m. salvini is separable from I. m. carrikeri by its larger bill (usually
over 16.5 mm from anterior margin of nostril to tip) and more extensive
black gular patch, although the latter is often considerably affected by the
‘‘make’’ of the skin. There is overlap in color characters; salvini seems
never to be orangish, as are many individuals of carrikeri, but the more
yellow examples of carrikeri could not be distinguished from salvini.

Fringillidae
Zonotrichia capensis

Although highly polytypic, Zonotrichia capensis shows an unusual lack
of differentiation from Costa Rica through Panama, the Andes of Colombia,
Ecuador, and western Venezuela. All specimens from these areas have in
the past been assigned to the subspecies Z. c. costaricensis Allen, 1891. On
the basis of nine specimens taken from Cerro Campana, western Panama
Province, Wetmore (1951) named a new subspecies, Z. c. orestera, distin-
guished from costaricensis by its supposedly darker coloration. In compar-
ing these birds with the extensive series in the USNM from Costa Rica,
Panama, and Colombia, I am unable to appreciate the distinction, however.
Because all specimens in the type series of orestera were collected within
a few days of each other in March, they are in a similar stage of molt and
wear, causing them to exhibit a certain uniformity in appearance that one
would not normally encounter in a more randomly selected series of spec-
imens. This may have influenced Wetmore’s conclusion. Nevertheless,
there are numerous examples of costaricensis in the USNM collections that

372 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

cannot be separated from those in the type series of orestera. I regard Z.
c. orestera Wetmore, 1951 as a synonym of Z. c. costaricensis Allen, 1891.
Considering the vast range through which Z. c. costaricensis shows no
appreciable variation, it would indeed have been remarkable if the birds
from Cerro Campana had differentiated.

Acknowledgments

I am indebted to the curators of the following institutions for lending
specimens or permitting their examination: American Museum of Natural
History, New York (AMNH); Academy of Natural Sciences of Philadelphia
(ANSP); Carnegie Museum of Natural History, Pittsburgh (CM); Field Mu-
seum of Natural History, Chicago (FM); Museum of Comparative Zoology,
Cambridge, Massachussets (MCZ); Museum of Vertebrate Zoology, Berke-
ley; National Museum of Natural History, Smithsonian Institution (USNM).
I am grateful to Eugene Eisenmann and Kenneth C. Parkes for providing
information and for their comments on the manuscript. This is contribution
number 8 of the Wetmore Papers, a project supported in part by trust funds
from the Smithsonian Institution for completing unfinished work and study
of undescribed material left by the late Alexander Wetmore.

Literature Cited

Bangs, O. 1902. On a second collection of birds made in Chiriqui, by W. W. Brown, Jr.—

Proceedings of the New England Zoological Club 3:15—70.

. 1906. Notes on birds from Costa Rica and Chiriqui, with descriptions of new forms

and new records for Costa Rica.—Proceedings of the Biological Society of Washington

19:101-112.

Blake, E. R. 1958. Birds of Volcan de Chiriqui, Panama.—Fieldiana: Zoology 36(5):499-577.

. 1968. Family Vireonidae. Family Icteridae. Jn R. A. Paynter, Jr., ed. Check-list of

Birds of the World. Volume 14.—Harvard University Press, Cambridge, Massachusetts.

Chapman, F. M. 1914. Diagnoses of apparently new Colombian birds, II.—Bulletin of the
American Museum of Natural History 33(12): 167-192.

Eisenmann, E. 1955. The species of Middle American birds.—Transactions of the Linnaean
Society of New York 7:vi + 128 pages.

Griscom, L. 1935. The ornithology of the Republic of Panama.—Bulletin of the Museum of
Comparative Zoology 78(3):261-—382.

Hellmayr, C. E. 1935. Catalogue of birds of the Americas. Part 8.—Field Museum of Natural

History Zoological Series 13(8):1—541.

. 1936. Catalogue of birds of the Americas. Part 9.—Field Museum of Natural History

Zoological Series 13(9):1—458.

. 1937. Catalogue of birds of the Americas. Part 10.—Field Museum of Natural History

Zoological Series 13(10): 1-228.

Kennard, F. H., and J. L. Peters. 1928. A collection of birds from the Almirante Bay region
of Panama.—Proceedings of the Boston Society of Natural History 38(10):443-465.

Meyer de Schauensee, R. 1951. The birds of the Republic of Colombia. Part 4.—Caldasia
SS.

VOLUME 94, NUMBER 2 37/3;

Nelson, E. W. 1912. Descriptions of new genera, species and subspecies of birds from Pan-
ama, Colombia, and Ecuador.—Smithsonian Miscellaneous Collections 60(3): 1-25.

Peters, J. L. 1931. Additional notes on the birds of the Almirante Bay region of Panama.—
Bulletin of the Museum of Comparative Zoology 71(5):293-345.

Sclater, P. L., and O. Salvin. 1869. Descriptions of three new species of tanagers from Ve-
ragua.—Proceedings of the Zoological Society of London 1869:439-440.

Slud, P. 1964. The birds of Costa Rica, distribution and ecology.—Bulletin of the American
Museum of Natural History 128:1—430.

Storer, R. W. 1970. Family Thraupidae. Jn R. A. Paynter, Jr., ed. Check-list of Birds of the
World. Volume 13.—Harvard University Press, Cambridge, Massachusetts.

Todd, W. E. C. 1917. Preliminary diagnoses of apparently new birds from Colombia and
Bolivia.—Proceedings of the Biological Society of Washington 30:3-6.

Wetmore, A. 1951. Additional forms of birds from Colombia and Panama.—Smithsonian
Miscellaneous Collections 117(2):1-11.

Zimmer, J. T. 1941. Studies of Peruvian birds. 39.—American Museum Novitates 1127:1-20.

. 1947. Studies of Peruvian birds. 52.—American Museum Novitates 1367:1-26.

National Museum of Natural History, Smithsonian Institution, Washing-
ton, D.C. 20560.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 374-377

SACCOGASTER MELANOMYCTER (OPHIDITFORMES:
BYTHITIDAE), A NEW FISH SPECIES
FROM THE CARIBBEAN

Daniel M. Cohen

Abstract.—The ophidiiform bythitid fish Saccogaster melanomycter, the
seventh known species of the genus, is described from a single, egg-bearing
female taken at 7.5 m from the Caribbean coast of Colombia near Santa
Marta. Other species have been trawled from 100-820 m. Differences from
its congeners include a black nasal chamber, fewer anal fin rays (37 versus
46-64), and fewer vertebral centra (45 versus 51-57). It is closest to S.
normae from Peru in having a partly restricted gill opening and in lacking
a elongate fleshy pectoral peduncle.

A single small bythitid fish collected from shallow water on the Caribbean
coast of Colombia and subsequently discovered in the Museum of Compar-
ative Zoology is described well by the generic diagnosis for Saccogaster,
a circumtropical genus of rarely encountered benthic fishes, presented by
Cohen and Nielsen (1972). However, this specimen is so different from the
six known species of Saccogaster that it cannot be identified with any of
them and is named below. Terminology and methods follow Cohen and
Nielsen (1972, 1978).

Saccogaster melanomycter, new species
Figs. 1, 2

Holotype.—The only known specimen, MCZ 47362, 2, Colombia, Carib-
bean, Santa Marta, Gairaca Bay (Ensenada de Gayraca), 25 feet, coral reef
environment; field no. RHCSM 77.

Diagnosis.—Nasal chambers sooty black, separated from the orbit by a
white area; anal fin rays 37 (46 to 64 in other species); vertebrae 45 (51 to
57 in other species); no elongate, fleshy, pectoral peduncle; dorsal extent
of gill opening at upper level of pectoral fin base; no scales on head or body.

Description.—Counts: dorsal 74; anal 37; pectoral 18; caudal 10; ventral
1-1; vertebrae 16 + 29 = 45; developed rakers on first arch 3. Measure-
ments in mm: SL 66 (approximate, due to distorted nature of specimen),
snout to dorsal fin origin 20.5, snout to anal fin origin 42.1, snout to ventral
fin base 15.0, body depth at vent 10.6, head length 18.2, snout length 3.4,
eye diameter 2.1, interorbital width 3.3, upper jaw length 9.5, maxilla depth

VOLUME 94, NUMBER 2 375

Fig. 1. Saccogaster melanomycter, holotype, MCZ 47362, 66 mm SL. Reconstructed view
by Keiko Hiratsuka Moore.

3.3, symphysis of cleithra to ventral fin 3.6, ventral fin length 7.4, pectoral
peduncle depth 4.8, pectoral fin length 7.1.

Body compressed, relatively short, greatest depth about 4.9 in SL. Pre-
anal about 1.6 in SL. Head compressed, about 3.6 in SL. Skin relatively
thick and opaque.

Lateral line continuous, originating above and anterior to upper angle of
gill opening and descending to midline slightly beyond midpoint of body; its
course marked by a narrow unpigmented line along which small papillae are
widely spaced, about a dozen darkly pigmented ones to level of vent, be-
yond which they are pale and difficult to count.

Lateral canal head pores 1, well above angle of gill opening; supraorbital

Fig. 2. Saccogaster melanomycter, holotype, MCZ 47362, 18.2 mm HL. Detailed view of
head by Keiko Hiratsuka Moore.

376 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

pores 1, on upper lip medial to the tubular anterior nostril; infraorbital pores
3, 2 on upper lip below posterior nostril, 1 above posterior section of max-
illary; mandibular pores 3, 2 large ones near tip of lower jaw, a small one
near the angular.

Snout fleshy, with loose skin, perhaps inflated in life. Jaws subequal.
Many small, dark filaments on the head, though absent from side of snout.
A short, strong opercular spine present. Upper rear part of maxillary free
and unsheathed.

Gill opening extending dorsally no farther than upper level of the pectoral
fin base.

First gill arch with 3 spiny protuberances that are only slightly larger than
the flat, spiny plates covering most of the arch. Gill filaments on first arch
neither notably long nor short, closest to those of S. maculata (Cohen and
Nielsen, 1972, fig. 1A). Pseudobranch of 2 filaments present.

Dentition on premaxillary a narrow, exterior band of granular teeth and
an inner single row of enlarged, needle-like teeth. Vomer with a broadly
U-shaped band of sharp-pointed teeth, the inner ones larger; a similar pat-
tern continues on the palatine. Dentary with an irregular exterior row of
small, slightly compressed teeth and an interior row of larger pointed teeth.

Dorsal fin originating over centrum 6, anal fin under 26 or 27. Adpressed
pectoral fin extending about one-third of distance from insertion of pectoral
fin rays to anus. Although the ossified pectoral radials are elongate as in
other Saccogaster species, they lie in the body wall and are not enclosed
in a long, free peduncle.

The type is a gravid female. The body cavity is nearly filled with two
distended ovaries packed with unfertilized eggs of about 1 mm diameter.
Pyloric caeca developed as two, small, slight hemispherical protuberances.

Color light brown, darker dorsally and on the caudal section. Dorsal and
anal fins with a narrow pale rim. Nasal capsule lined with sooty black pig-
ment, about equal in size to orbit; area between the eye and capsule im-
maculate white.

Abdominal centra 16; neural spines on the middle ones variously de-
pressed and truncate. First parapophyses on centrum 6.

Habitat.—The capture of S. melanomycter at a depth of about 7.5 m in
a reef area is surprising, as all previously reported examples have been
trawled at depths ranging from 100 to 820 m.

Relationships.—Saccogaster melanomycter falls neatly into the scheme
of possible relationships for species of Saccogaster proposed by Cohen and
Nielsen (1972). Although §. melanomycter is readily distinguished from all
species of the genus by its low anal fin and vertebral counts and its distinc-
tive nasal pigmentation pattern, it fits in the parva-tuberculata-normae
group of species because of its continuous lateral line and lack of scales.
Within this group, S. melanomycter shares two specialized characters with

VOLUME 94, NUMBER 2 ST

S. normae, a restricted gill opening and absence of a free pectoral peduncle
for the elongate pectoral radials. Saccogaster melanomycter is, however,
less specialized than S. normae in having a larger gill opening (although
smaller than in other Saccogaster species) and in having the rear of the
maxillary free.

Etymology.—The species name is taken from the Greek melano-, black,
and mykter, nose.

Acknowledgments

i am grateful to Karsten Hartel for allowing me to Study the specimen
described in this paper and to Bruce Collette and J grgen Nielsen for reading
the manuscript.

Literature Cited

Cohen, D. M., and J. G. Nielsen. 1972. A review of the viviparous ophidioid fishes of the
genus Saccogaster.—Proc. Biol. Soc. Wash. 85:445—-468.

, and . 1978. Guide to the identification of genera of the fish order Ophidiiformes
with a tentative classification of the order—NOAA (Natl. Ocean. Atmos. Adm.) Tech.
Rep. NMFS (Natl. Mar. Fish. Serv.) Circ. 417, 72 pp.

Systematics Laboratory, National Marine Fisheries Service—NOAA,
National Museum of Natural History, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 378-379

FALSIPATELLINA, A NEW NAME FOR
PSEUDOPATELLINA KENAWY AND
NYIRO, 1967 NON HAQUE, 1960
(FORAMINIFERIDA)

Drew Haman and Richard W. Huddleston

Haque (1960:28) established the genus Pseudopatellina (type-species P.
arthurcooperi n. sp.) for specimens obtained from the middle to late Eocene
shales underlying the Khirthar limestone at Ush Bul spring, Sor Range,
West Pakistan. This group has been recorded in some systematic compila-
tions (Tobias et al., 1970) but is absent from a number of leading works on
foraminiferid systematics (Loeblich and Tappan, 1964a, b; 1974). Conse-
quently, this genus is not well known.

Kenawy and Nyiro (1967:104) used the generic epithet Pseudopatellina
(type-species P. plana n. sp.) for specimens obtained from upper Oligocene
Strata at Eger, Hungary.

Pseudopatellina Kenawy and Nyiro, 1967, is a junior homonym of Pseu-
dopatellina Haque, 1960, and in accordance with ICZN Art. 53 is herein
renamed Falsipatellina.

Acknowledgments

The authors are indebted to Prof. Dr. Abbas Ibrahim Kenawy, University
of Assiut, Egypt, U.A.R., for allowing them to effect this name change, and
to Chevron Oil Field Research Company for publication permission.

Literature Cited

Haque, A. F. M. M. 1960. Some middle to late Eocene smaller Foraminifera from the Sor
Range, Quetta District, West Pakistan.—Pakistan Geological Survey Memoirs, Palaeon-
tologia Pakistanica 2(2):1-79.

Kenawy, A. I., and R. M. Nyird. 1967. Zwei neue Foraminiferen aus dem Oberoligozan in
Eger (Nordungarn).—Annales Historico-Naturales Musei Nationalis Hungarici, Pars
Mineralogica et Palaeontologica 59:103-107.

Loeblich, A. R. Jr., and H. Tappan. 1964a. Sarcodina, chiefly ‘“Thecamoebians”’ and Fora-

miniferida. Pp. Cl—C900 in R. C. Moore, ed., Treatise on Invertebrate Paleontology,

Part C, Protista 2.—Kansas University Press, xxxi + 900 pp.

, and . 1964b. Foraminiferal classification and evolution.—Journal of the Geo-

logical Society of India 5:5—40.

VOLUME 94, NUMBER 2 379

, and . 1974. Recent advances in the classification of the Foraminiferida. Pp. 1-
53 in R. H. Hedley, and C. G. Adams, Foraminifera; vol. 1.—Academic Press, viii +
276 pp.

Tobias, M., E. Lilley, and M. Oxenham. 1970. Fossil Protozoa.—The Zoological Record,
1967, vol. 104, sec. 2, Protozoa, pt. II: 145-198.

Chevron Oil Field Research Company, P.O. Box 446, La Habra, Califor-
nia 90631.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 380-390

THE NATURE OF THE VARIABILITY IN THE
VARIABLE SEEDEATER IN PANAMA
(SPOROPHILA AMERICANA: EMBERIZINAB)

Storrs L. Olson

Abstract.—Detailed examination of variation in Sporophila americana in
Panama necessitates major changes in the stated ranges and nomenclature
of the subspecies found there. The name hicksii (Lawrence, 1865) is revived
for white-throated black and white birds on the Pacific coast of Panama
from Veraguas east through Darién (skipping the Canal Zone), the Carib-
bean coast in San Blas, and south in western Colombia to Valle. The cur-
rently recognized name chocoana (Meyer de Schauensee, 1950) becomes
a synonym of hicksii. It is suggested that the type locality of hicksii should
be Buenaventura, Colombia, instead of Panama. Black-throated black and
white birds from the Pacific slope of Costa Rica and Chiriqui may now be
known under the resurrected name hoffmanni Cabanis, 1861. The nearly
black subspecies corvina (Sclater, 1859) extends along the western Carib-
bean coast of Panama and in the Canal Zone intergrades with hicksii to form
a hybrid zone in which no pure parental types occur. The long-used but
dubiously applicable name “‘aurita’’ (Bonaparte, 1850) is tentatively applied
to these intergrades and the specimens are analyzed by the use of a hybrid
index.

As its name implies, the Variable Seedeater, Sporophila americana, has
been regarded as being so variable in the adult male plumage, particularly
in Panama, as to defy interpretation in terms of normal geographic variation.
This idea became so fixed in the earlier literature that contrary information
was often ignored. No recent attempts have been made to examine series
of specimens from Panama to elucidate distributional patterns there. By
current taxonomic practices there are three subspecies of Sporophila amer-
icana recognized in Panama: S. a. corvina, a nearly all black form found
on the Carribean slope of western Panama north to Mexico; S. a. aurita,
a supposedly extremely variable black and white form found on the Pacific
slope of Costa Rica and Panama to the Canal Zone and on both slopes east
of the Canal to Darién, where it is supposedly replaced by a more constant
white-throated form, S. a. chocoana, inhabiting easternmost Panama and
Pacific Colombia south to the Rio Dagua. The development of this classi-
fication may be traced through the publications of Sclater (1871), Chapman

VOLUME 94, NUMBER 2 381

- corvina °
- almost pure corvina O
-hicksii wees

hoffmanni

-hoffmanni x hicksii intergrades cya

-"aurita" (hybrid corvina x hicksii)

x O 80068

Fig. 1. Distribution of the forms of Sporophila americana in Panama, based on specimens
examined in this study.

(1926), Hellmayr (1938), Meyer de Schauensee (1950, 1952) and Eisenmann
(1957).

Not all of the observations below are entirely original, but their signifi-
cance has not hitherto been appreciated. I shall not attempt to treat the
history of each conclusion, as this may be found in the references just cited.
The classification I shall propose differs considerably from that above. For
this reason and because the nomenclatural changes I shall propose are rather
intricate, I shall begin by presenting my conclusions first and documenting
them beyond.

The revised distribution and nomenclature of S$. americana in Panama is
shown in Fig. 1. No change in status is required for the black populations
from the western Caribbean coast, S. a. corvina. Much of the rest of the
isthmus is occupied by a black and white subspecies, with a white rump and
throat, which occurs in most of Panama east of the Canal Zone and west of
the Canal Zone on the Pacific slope to central Veraguas. The name hicksii
is resurrected for this subspecies, with chocoana falling into synonymy. A
similar subspecies, but with a black throat, is found from western Veraguas,
including Isla Coiba, west along the Pacific slope through Costa Rica. The
name hoffmanni is revived for this form. The only birds showing great
variability are restricted to the area of the Canal Zone, where populations
consist entirely of intergrades between corvina and hicksii, and where no
pure parental types occur. The name ‘‘aurita,’’ the application of which is
dubious in any case, is tentatively used for these intergrades.

382 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Sporophila americana corvina (Sclater)

Spermophila corvina Sclater, 1859, Proc. Zool. Soc. London 1859:379.
Playa Vicente, Oaxaca, Mexico.

Spermophila badiiventris Lawrence, 1865, Ann. Lyc. Nat. Hist. N.Y. 8:172.
Greytown, Nicaragua. Type re-examined for this study.

Characters.—Adult males almost entirely black, with only the speculum
of the wing and midline of belly white. Speculum smaller than in hicksii.
Females and subadult males are markedly darker and more sooty than in
other subspecies, but have not been considered in determining the distri-
bution or extent of intergradation in any of the forms.

Range.—Caribbean slope of southeastern Mexico, south through Bocas
del Toro, Panama. The few specimens known from the Caribbean slope
between Bocas del Toro and the Canal Zone are nearly pure corvina, but
show faint traces of white on the sides of the neck. I have designated these
as ‘‘0+’’ in the hybrid index discussed in the account of S. a. “‘aurita,”’
but have listed these specimens below under corvina.

Specimens examined (only specimens from Costa Rica and Panama are
listed although others were examined).—COSTA RICA. GUANACASTE:
ca. 3 miles E Tilaran, 950 m (1, LSU); Arenal, 500 m (1, LSU). ALA-
JUELA: Naranjo (1, USNM); Villa Quesada (1, USNM). HEREDIA: Puer-
to Viejo (1, LSU). CARTAGO: Bonilla (6, USNM; 4, AMNH); Guayabo
(3, USNM); Tucurrique (1, AMNH); Aquiares (1, AMNH). SAN JOSE:
Carillo (3, AMNH; 1, USNM). LIMON: Cariari (4, WFVZ); Finca La Lola,
Rio Madre de Dios (1, WFVZ); Jiménez (2, USNM); San Bernardo (1,
USNM); Sipurio (2, USNM); Uva (1, USNM); Limon (3, AMNH); Guapiles
(2, AMNH); Siquirres (1, AMNH); Atalanta (2, AMNH).

PANAMA. BOCAS DEL TORO: Almirante (5, USNM; 4, AMNH);
Changuinola (1, USNM); Cocoplum (7, ANMH); “**‘Bocas del Toro’’ (1,
ANSP). The following were ranked °*‘0+,”’ or almost pure corvina, in the
hybrid index discussed below: VERAGUAS: Rio Calovévora, Caribbean
slope (1, AMNH); COCLE: Tigre, head of Rio Guabal (1, USNM); El Ur-
acillo, Rio Negro (1, USNM). COLON: Chilar, Rio Indio (1, USNM). CA-
NAL ZONE: Gatun (1, USNM); Gamboa Pipeline Road (1, USNM).

Sporophila americana hicksii (Lawrence)

Spermophila hicksii Lawrence, 1865, Ann. Lyc. Nat. Hist. N.Y. 8:171.
‘‘Panama’’ (probably = Buenaventura, Colombia, see below). Type re-
examined for this study.

Spermophila aurita chocoana Meyer de Schauensee, 1950, Proc. Acad. Nat.
Sci. Phila. 52:138, Nuqui, Choco, Colombia.

VOLUME 94, NUMBER 2 383

Characters.—Adult males with throat, sides of neck, lower breast and
abdomen, rump, undertail coverts, and speculum white; pectoral band and
variable amount of chin black.

Range.—Pacific coast of Panama from central Veraguas and the Azuero
Peninsula east (skipping the Canal Zone) through Panama Province and
Darien, the Atlantic slope in San Blas, south along the Pacific slope of
Colombia to the vicinity of the Rio Dagua, Valle.

Specimens examined.—PANAMA. VERAGUAS: Sona (5, USNM); La
Colorada, Santiago (2, AMNH); E shore of Montijo Bay, | mile S of Angulo
River mouth (2, CM); Isla Gobernadora (2, USNM; one of these is more
similar to those from Isla Coiba). LOS SANTOS: Tonosi (1, USNM). CO-
CLE: Gago (1, USNM). PANAMA PROVINCE: Pacora (1, USNM); Chi-
man, Rio Chiman (1, USNM); Charco del Toro, Rio Majé (2, USNM). SAN
BLAS: Mandinga (2, USNM); Permé (2, MCZ); Puerto Obaldia (3, USNM,
1 MCZ). DARIEN: Cana (4, USNM; 3, MCZ); mouth of Rio Paya, Rio
Tuira (1, USNM); Pucro, Rio Pucro (1, USNM); Jaqué (5, USNM); El Real,
Rio Tuira (5, AMNH); Boca de Cupe, Rio Tuira (3, AMNH); Rio Sambu
(1, ANSP).

COLOMBIA. CHOCO: Nuqui, Rio Jurubida (4, ANSP); Nuqui (2,
USNM); Acandi (5, USNM); Jurado (1, AMNH; 1, ANSP); Rio Baudonde
(1, ANSP); Quibdo (1, ANSP); upper Rio Baudo (1, ANSP); Andagoya (1,
ANSP). ANTIOQUIA: Dabeiba, Rio Sucio (1, AMNH); Villa Arteaga (2,
USNM). CORDOBA: Socarré, Rio Sinu (1, USNM). CALDAS: Santa Ce-
cilia (3, ANSP). VALLE: Punta Muchimbo, Rio San Juan, | mile S of mouth
of Rio Calima (2, USNM); San José (1, USNM); Buenaventura (2, USNM).

Remarks.—When Meyer de Schauensee (1950) described chocoana, he
compared it with birds from Costa Rica, western Panama, and the Canal
Zone, all of which were considered to represent aurita and all of which
have black throats. He recognized that birds from Colombia and Darién
differed in consistently possessing a white throat and he therefore segre-
gated these under the name chocoana. A major problem arises here in the
nature and orgin of the type specimen of hicksii Lawrence (USNM 40300).
Meyer de Schauensee (1950:139) reported that “‘Mr. Herbert Deignan has
kindly examined the type of hicksii for me and writes me that although it
is a white-throated example, it was collected by Hicks ‘probably not far
from the city of Panama.’ It therefore must be regarded as an aberrant
example of S. a. aurita.”’

The type of hicksii, which I have examined, is a perfectly typical example
of chocoana, with a pure white throat and rump. It shows not a trace of
the intergradation with corvina that marks ‘‘aurita.’’ The specimen is la-
belled simply ‘““Panama,”’ the implication being that it came from the vicinity

384 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

of Panama City, on the Pacific slope. No such individuals with pure white
throats from the vicinity of Panama City or the Canal Zone exist in any of
the collections I have examined, although they have been taken on the
Pacific slope both to the east and to the west of the Canal Zone area.

The type of hicksii was sent to S. F. Baird at the Smithsonian Institution
by Fred Hicks in a lot of 54 specimens, all of which came from Panama
except for 4 from Cali, Colombia, and 12 from Buenaventura, Colombia
(letter of 27 December 1864 from Hicks to Baird; accession file 588, USNM).
Not all of these specimens were catalogued. The type of hicksii was cata-
logued immediately following 3 specimens of the same species labelled as
being from Buenaventura, one of which (USNM 40299) is now missing.
These are the only other specimens of Sporophila americana among those
collected by Hicks; the type of hicksii is the only one labelled as coming
from Panama.

Hicks’ original labels were for the most part more informative than usual
for the period, and often included soft part colors and measurements, as
well as locality, date, collector, and field number. Unfortunately, these were
made of very fragile paper and in many instances appear to have been
deliberately removed from the specimens, as is the case with the type of
hicksii. Whoever copied the data onto the new labels often did not transfer
all of the information that was on the originals and the transcriptions were
also not without error. There is a specimen of the tanager Ramphocelus
flammigerus icteronotus in the same lot as the type of hicksii that has **Pan-
ama’’ written on one side of the label and ‘‘Buenaventura’’ on the other.

Circumstantial evidence strongly suggests that the type specimen of Sper-
mophila hicksii Lawrence came from Buenaventura, Colombia; it is unlikely
to have been obtained in the vicinity of Panama City. Even if the specimen
did come from Panama, it can not be regarded as an abnormal specimen of
‘‘aurita,’’ since similar white-throated populations occur on the Pacific slope
of Panama both to the west and to the east of the Panama City—Canal Zone
area. The name Spermophila aurita chocoana Meyer de Schauensee, 1950
therefore becomes a junior synonym of Spermophila hicksii Lawrence, 1865
and I recommend that Buenaventura, Valle, Colombia, be regarded as the
type locality of hicksii.

Buenaventura is at or near the southern limit of the range of hicksii.
Meyer de Schauensee (1950) correctly recognized the Rio Dagua as the
approximate line of demarcation between chocoana (=hicksii) and the sub-
species ophthalmica (Sclater, 1860; type locality Babahoyo, Ecuador),
which differs in having a narrower pectoral band and whiter chin. Colombian
birds from Los Cisneros, Valle (3, AMNH), Guapi, Cauca (1, USNM), and
from Narino through Ecuador are referable to ophthalmica.

The discovery of white-throated individuals of hicksii west of the Canal
Zone is significant (see discussion of “‘aurita’’). Although the amount of

VOLUME 94, NUMBER 2 385

black in the chin is variable in hicksii, the birds from Coclé and the Azuero
Peninsula stand out in having the chin almost totally white, thus contrasting
markedly with the intergrades (‘‘aurita’’) to the east and with the subspecies
hoffmanni to the west.

Sporophila americana hoffmanni Cabanis

Sporophila hoffmanni Cabanis, 1861, Journ. Orn. 9:6. Costa Rica.

Spermophila collaris Lawrence, 1865, Ann. Lyc. Nat. Hist. N.Y. 8:177.
David, Chiriqui, Panama. (Not Loxia collaris Boddaert.) Type re-exam-
ined for this study.

Characters.—Similar to hicksii but adult males with throat black, with
varying amounts of white, either as a narrow collar or as discontinuous
patches, on either side of the neck.

Range.—Pacific slope of Costa Rica and Panama in Chiriqui and Isla
Coiba, intergrading with hicksii in western Veraguas.

Specimens examined.—COSTA RICA. SAN JOSE: El General (7, MCZ).
PUNTARENAS: Tambor, Nicoya (3, LACM); Las Agujas (1, LACM);
Pigres (1, USNM); Pozo Azul (5, CM; 1, MCZ); El Pozo del Rio Grande
(de Térraba) (3, MCZ; 4, CM; 2, FM; 1, ANSP); Buenos Aires (3, CM; 4,
FM; 4, AMNH); Puerto Jiménez, Peninsula de Osa (1, CM; 1, AMNH);
Rincon de Osa (1, LSU; 1, WFVZ); Boruca (1, ANSP; 1, AMNH; 2, CM;
1, USNM; 1, ANSP; 7, MCZ); 13 km S of Palmar Sur (3, WFVZ); Hele-
chales (7, WFVZ).

PANAMA. CHIRIQUI: David (1, USNM; 1, MCZ); Divala (1, MCZ; 1,
USNM); El Volcan (4, USNM); Buena Vista (1, USNM); Puerto Armuelles
(4, USNM); Boqueron (1, AMNH). VERAGUAS: Isla Coiba (7, USNM;
4, AMNH [collected by Batty, locality not reliable]).

Remarks.—Although Chapman (1926) recognized this subspecies (under
the name collaris) and Hellmayr (1938) conceded that birds from Costa Rica
and Chiriqui were generally separable from ‘‘aurita,’’ this western subspe-
cies is not at present recognized, due to its similarity to certain of the
intergrades from the Canal Zone and the fact that, hitherto, hicksii was not
known to occupy the intervening area. Some of the intergrades of ‘‘aurita”’
are indeed very similar to the western birds, but they may usually be dis-
tinguished by the lesser amount of white in the rump. Within the populations
from the Pacific slope of Costa Rica and Chiriqu1, there is little variation in
adult male plumage. Yet because of the confusion that has long attended
the concept of variation in this species, Wetmore (1957) assigned a series
of seven males from Isla Coiba, which closely resemble one another and
show little variation, to the subspecies aurita, which was then considered
to be extremely variable.

The earliest available name for the black-throated western birds is hoff-

386 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

manni Cabanis, 1861, the original description of which clearly pertains to
this population. S$. a. hoffmanni evidently intergrades with S. a. hicksii in
Veraguas. A series of five males from Sona are closest to hicksii but have
more black in the chin than typical of most individuals of that subspecies.
The birds from Isla Coiba are like hoffmanni but show a slight tendency
towards hicksii in having the white collar mostly continuous and a few white
feathers on the chin. Of two adult males from Isla Gobernadora, Veraguas,
one resembles the birds from Isla Coiba whereas the other could pass as
typical hicksii.

Sporophila americana *‘aurita’’ (Bonaparte)

Spermophila aurita Bonaparte, 1850, Consp. Gen. Av. 1(2):497 ‘‘Bras.
[il]? = Panama?

Spermophila semicollaris Lawrence, 1863, Ann. Lyc. Nat. Hist. N.Y. 8:10.
Lion Hill, Panama Railroad. Type re-examined for this study.

Spermophila fortipes Lawrence, 1865, Ann. Lyc. Nat. Hist. N.Y. 8:172.
Line of Panama Railroad. Type re-examined for this study.

Characters. —Extremely variable, ranging from individuals similar to
hicksii, but with slightly more black in the rump, throat, or belly, to indi-
viduals nearly as black as corvina; see discussion of hybrid index below.

Range.—Essentially restricted to the Canal Zone area of central Panama;
known as far west as La Chorrera, western Panama Province, and as far
east as Buenaventura Island, near Portobelo, Colon, and the Candelaria and
Peluca Hydrographic Stations at the head of Madden Lake.

Specimens examined.—All are from the Canal Zone or immediately ad-
jacent parts of Panama or Colon provinces, except where indicated, and all
are from the USNM, AMNH, and MCZ collections except 3 from Gatun
(ANSP) and 1 from Barro Colorado Island (FM). Hybrid index (HI) is given
first, followed by number of specimens in parentheses. Aspinwall, HI-1 (1);
Balboa, HI-3 (2); Barro Colorado Island, HI-5 (1); Bas Obispo, HI-1 (1);
Bohio HI-4 (1); Buenaventura Island, near Portobelo, Colon Province, HI-
3 (1); Candelaria Hydrographic Station, HI-3 (3); Chiva Chiva, HI-4 (1);
Colon, HI-1 (1), HI-2 (1); Corozal, HI-3 (2); Curundu HI-S5 (1); Fort Loren-
zo, HI-1 (1); Frijoles, HI-1 (1), HI-4 (1); Gamboa, HI-2 (1); Gatun, HI-1 (5),
HI-2 (7), HI-3 (4), HI-4 (1); Juan Mina, HI-1 (1), HI-3 (2), HI-4 (1), HI-5
(2); La Chorrera, western Panama Province, HI-2 (1); Lion Hill (Loma del
Leon), HI-1 (1), HI-2 (4); HI-3 (3); HI-4 (2); Miraflores, HI-3 (1), HI-5 (1);
line of Panama Railroad, HI-5 (1); ‘“‘near Panama,’’ HI-2 (2), HI-3 (4), HI-
4 (1), HI-5 (1); “*Panama,’’ HI-2 (3); Peluca Hydrographic Station, HI-3 (1);
Tabernilla, HI-2 (1), HI-3 (1), HI-4 (1); Rio Trinidad, Agua Clara, HI-2 (1);
Savanna, HI-3 (1).

VOLUME 94, NUMBER 2 387

Remarks.—There is considerable doubt in my mind about the proper use
of the name aurita Bonaparte, long applied to most of the specimens of this
species from Panama and Pacific Costa Rica. Bonaparte’s (1850:497) entire
original description reads as follows: ‘‘Sp. aurita, Bp. Mus. Paris. ex Bras.
Similis praecedenti [Sporophila luctuosa]; gula pectoreque nigerrimis: sed
rostro nigro et macula utrinque auriculi alba.’’ This description could apply
to many individuals from the Canal Zone or equally well to hoffmanni of
Costa Rica and Chiriqui. Sclater (1871:15) considered the specimens he
examined from ‘‘Panama and Chiriqui’’ to represent a single species and
stated that “‘the typical example of S. aurita in Mus. Paris clearly belongs
to it, though the locality (Brazil) attributed to it in the ‘Conspectus’ is no
doubt erroneous.’’ It is not absolutely clear that Sclater actually examined
the type; Hellmayr (1938:191) could not find it at the Paris Museum in 1935.
Because of Sclater’s action, the type locality of aurita has since been taken
as Panama, but neither the true identity nor the provenance of the type has
been positively determined.

Several nomenclatural resolutions suggest themselves. One could regard
the name aurita Bonaparte as being of undeterminable application, or one
could go on the assumption that the name was correctly applied to birds
from the hybrid zone in the central part of the Panamanian isthmus. In the
latter case, one might attempt to apply the name aurita, which is the earliest
available for any of the Panamanian subspecies, to one or the other of the
parental phenotypes, i.e. either to corvina or hicksii. To do this success-
fully, one would still need the holotype of aurita to determine to which
parental type it was most similar. Were it to prove more similar to the black
subspecies, then corvina would become a synonym of aurita, thus causing
endless confusion, since corvina is the one name that has never been applied
to the more variable populations in Panama. I have adopted a compromising
course by applying the name ‘‘aurita,’’ in quotes, only to those individuals
that are clearly intergrades between corvina and hicksii. Although this is
not altogether a satisfactory resolution, this population has a circumscribed
range in which neither parental phenotype occurs and in which virtually all
individuals can be certainly identified as belonging to “‘aurita,’’ as opposed
to some other population. This also prevents the sudden disappearance of
the name under which the majority of Panamanian birds have appeared in
most of the earlier literature.

To facilitate analysis of variation in “‘aurita,’’ I grouped adult male spec-
imens according to the following hybrid index (HI).

HI 0.—Pure corvina. All black except for small white speculum and
white along midline of belly. Individuals fitting this description but
having faint traces of white ‘‘ears’’ I designated as ‘‘0+,’’ shown in
Fig. 1 as “‘almost pure corvina.”’

388 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON |

HI 1.—More white in midline, particularly lower abdomen; some white
in crissum. No, or almost no, white in rump. White “‘ears’’ present,
but sometimes vestigial.

HI 2.—Like HI 1 but more white in belly and crissum and light tips to
feathers on flanks away from midline. A few white-tipped feathers on
rump.

HI 3.—White pattern of abdomen evident but suffused with black. White
‘‘ears’’ extending onto throat as a variable collar. Some white on chin
and a little more on rump.

HI 4.—Belly mostly whitish, rump considerably white, but little white on
chin.

HI 5.—Like hicksii but less white in rump. }

HI 6.—Pure hicksii; black with white throat, white ‘‘ears,’’ white lower
breast and abdomen, white undertail coverts, speculum, and rump
(but not upper tail coverts), chin variably black, pectoral band black.

Disregarding the few “*0+’’ specimens from outside the area of the Canal
Zone, I examined 80 specimens of ‘‘aurita.’’ The number in each HI cate-
gory was as follows:

HI 0+ 2 3%)
HI 1 12 (15%)
HI 2 24 (30%)
HI 3 26 (33%)
HI 4 9 (11%)
HI 5 7 (9%)

Thus, 63% of the specimens could be regarded as more or less intermediate
between the two parental types, 18% were more like corvina and 20% were
more like hicksii. No specimens of pure parental stock occur in the area of
intergradation, and this population thus constitutes a hybrid zone by Short’s
(1969) definition.

Eisenmann (1957:260) came close to a correct interpretation in hypothe-
sizing that the variability of this species in Panama might be the result of
‘interbreeding between aurita and corvina’’ and that “‘nominate aurita is
itself the product of some earlier contact between corvina and a population
essentially like ophthalmica.’’ It was not apparent at that time, however,
that “‘aurita’’ was confined entirely to the Canal Zone area, nor that the
‘“population essentially like ophthalmica,’’ in the form of hicksii, still exists
in Panama on the Pacific slope on either side of the hybrid “‘aurita’’ zone.
Because the range of hicksii was obviously once continuous across the
Pacific side of the isthmus, it is evident that the new development is the
introgression by corvina genetic stock, thus creating a hybrid zone that has
fragmented the distribution of hicksii.

VOLUME 94, NUMBER 2 . 389

The age of the establishment of the hybrid zone between hicksii and
corvina is not certain, but Eisenmann’s (1957:260) supposition that the
forms in question “‘were probably not in contact’? a century ago, may be
doubted. The earliest specimens from the area of the Canal Zone are the
types of semicollaris Lawrence and fortipes Lawrence, collected by
McLeannen along the Panama Railroad in the mid-1800’s. I have examined
these and would classify both as HI 2 intergrades, with the type of fortipes
having less white on the rump and more white on the chin than the type of
semicollaris. Two other specimens collected by McLeannen I would clas-
sify as HI 2 and HI 5. Although I analyzed all the specimens of ‘‘aurita’’
from the hybrid zone by date and locality, I could detect no pattern of
change through time. This may be due to lack of adequate samples taken
in the 19th century. The hybrid zone has apparently been in existence for
Over a century and appears to be stable. The influence of corvina has not
yet spread much beyond the Canal Zone on the Pacific slope. On the other
hand, there is evidence, although slight, that the influence of hicksii may be
spreading westward along the Atlantic slope.

Acknowledgments

I am grateful to the curators and staff of the following institutions for
lending specimens or permitting their examination: American Museum of
Natural History, New York (AMNH); Academy of Natural Sciences of
Philadelphia (ANSP); Carnegie Museum of Natural History, Pittsburgh
(CM); Field Museum of Natural History, Chicago (FM); Natural History
Museum of Los Angeles County, Los Angeles (LACM); Museum of Zool-
ogy, Louisiana State University, Baton Rouge (LSU); Museum of Compar-
ative Zoology, Harvard University, Cambridge, Massachusetts (MCZ);
Western Foundation of Vertebrate Zoology, Los Angeles (WFVZ); Na-
tional Museum of Natural History, Smithsonian Institution, Washington,
D.C. (USNM). The figure is by Sigrid K. James. This is contribution number
9 of the Wetmore Papers, a project supported in part by trust funds from
the Smithsonian Institution for completing unfinished work and study of
undescribed material left by the late Alexander Wetmore.

Literature Cited

Bonaparte, C. L. 1850. Conspectus Generum Avium. Part 1. E. J. Brill, Leyden.

Chapman, F. M. 1926. The distribution of bird-life in Ecuador.—Bulletin of the American
Museum of Natural History 55:1-784.

Eisenmann, E. 1957. Notes on the birds of the Province of Bocas del Toro, Panama.—Condor
59(4):247-262.

Hellmayr, C. E. 1938. Catalogue of birds of the Americas. Part 11.—Field Museum of Natural
History Zoological Series 13(11): 1-662.

390 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Meyer de Schauensee, R. 1950. Colombian zoological survey. Part VII. A collection of birds

from Bolivar, Colombia.—Proceedings of the Academy of Natural Sciences of Phila-

delphia 102:111—139.

. 1952. A review of the genus Sporophila.—Proceedings of the Academy of Natural

Sciences of Philadelphia 104: 153-196.

Sclater, P. L. 1871. A revision of the species of the fringilline genus Spermophila.—Ibis,
series 3, 1(1):1-23.

Short, L. L. 1969. Taxonomic aspects of avian hybridization.—Auk 86(1):84—105.

Wetmore, A. 1957. The birds of Isla Coiba, Panama.—Smithsonian Miscellaneous Collections
134(9): 1-105.

National Museum of Natural History, Smithsonian Institution, Washing-
ton, D.C. 20560.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 391-403

WESTERN ATLANTIC SEA CUCUMBERS OF THE
GENUS THYONE, WITH DESCRIPTION OF TWO
NEW SPECIES (ECHINODERMATA: HOLOTHUROIDEA)

David L. Pawson and John E. Miller

Abstract.—Current status of each of the several Western Atlantic holo-
thurians referred to the genus Thyone s.l. is discussed. Two new species,
Thyone adinopoda and T. crassidisca, from the southeastern United States
are described.

Dendrochirotid holothurians of the genus Thyone s.l. are burrowing,
U-shaped forms with tube feet scattered over the body wall and not re-
stricted to the ambulacral radii. They usually occur in muddy to sandy
habitats, and range from the subtidal zone to depths in excess of 200 meters.
Systematic study of ‘“Thyone”’ species is rendered difficult because in many
species the diagnostic characters may change with growth, and in many the
calcareous ossicles of the body wall tend to disappear with growth.

Thyone currently comprises a miscellany of approximately 40 species. Of
the 14 Western Atlantic species referred to Thyone by Deichmann (1930),
only 2 remain in that genus today. The other species are now scattered
among several genera in 2 dendrochirotid subfamilies. Panning (1949, and
later papers) has been largely responsible for the reassignment of these
species. In her later papers, Deichmann (1954, 1957, 1963) ignored Panning’s
revision of the Cucumariidae, and attempted some revisions independently,
thus further confusing an already complex situation.

In this paper we attempt to bring up-to-date the nomenclature of the
Western Atlantic ‘““Thyones’’ by listing the currently accepted names for all
species. In addition, in the course of a monographic study of Western At-
lantic holothurians, we have found two new species of Thyone s.s., and the
opportunity is taken to describe these here.

Type-material is deposited at the National Museum of Natural History,
Smithsonian Institution (USNM), the Indian River Coastal Zone Museum,
Harbor Branch Foundation-Smithsonian Institution, Fort Pierce, Florida
(IRCZM), and the Florida Department of Natural Resources, St. Petersburg,
Florida (FSBC). This paper is Harbor Branch Foundation, Inc. Contribution
to Science No. 199. Financial support for this study was received from the
Harbor Branch Foundation, Inc. and from the Smithsonian Institution Fort
Pierce Bureau.

We thank Dr. Thomas S. Hopkins for allowing one of us (DLP) to study

392 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

holothurians collected under the auspices of the U.S. Bureau of Land Man-
agement, and the Florida Department of Natural Resources for giving us
access to holothurians collected by the ‘‘Hourglass’’ Cruises and other in-
vestigations in the Gulf of Mexico.

Current Concept of the Genus Thyone

When Panning (1949:468) transferred species from Havelockia sensu Pan-
ning, 1949 to Thyone sensu Panning, 1949, he suggested that Thyone in-
cludes 2 groups of species; Group I comprising those species with tables
but no rosettes in the introvert, and Group II comprising species with ro-
settes but no tables in the introvert. At least 3 Western Atlantic species,
pseudofusus Deichmann, inermis Heller, and crassidisca n. sp. fall into yet
another category, a Group III, where the introvert contains rosettes and
tables.

A thorough revision of the “‘supergenus’’ 7hyone is urgently needed, but
is beyond the scope of this paper. While some species are well-defined,
others appear to change greatly with growth, so that adults and young spec-
imens may be referred to different species. Deichmann (1947) tended to
dismiss Madsen’s (1941) careful analysis of 7. fusus, and his descriptions
of new taxa. While Madsen’s new species might not be valid, his detailed
descriptions are excellent, and should form a model upon which a thorough
revision of Thyone can be based.

Western Atlantic ‘“Thyone’’ Species Listed or Described by
Deichmann (1930, 1946)

Thyone briareus (Lesueur, 1824). Panning (1949) transferred this species
to Sclerodactyla.

Thyone scabra Verrill, 1873. Panning (1949) transferred this species to
Havelockia.

Thyone fusus (Miller, 1776). Deichmann (1930) described specimens of
Thyone which she found to be very closely similar to T. fusus (Miller) from
Northern Europe. In later papers, Deichmann (1946, 1947, 1954) concluded
that the West Indian “‘fusus’’ and the Mediterranean ‘‘fusus’’ were identical,
and that they differed in some respects from the typical northern European
T. fusus (Muller). Deichmann applied the name T7. inermis Heller to the
Mediterranean-W. Atlantic species, and this name is accepted here.

Thyone pseudofusus Deichmann, 1930. Unchanged.

Thyone belli Ludwig, 1887. Panning (1949) transferred this species to
Pseudothyone. Deichmann (1954) referred it to her (1941) genus Neothyone.
Panning’s action appears to be the more logical.

Thyone micropunctata Sluiter, 1910. Deichmann (1954) showed that this
species is a junior subjective synonym of Thyone belli Ludwig, 1887.

VOLUME 94, NUMBER 2 393

Fig. 1. Left, Ti hyone adinopoda, holotype, total length 26 mm; Right, Thyone crassidisca:
upper, holotype, total length 51 mm; lower. paratype, total length 40 mm.

Thyone cognata (Lampert, 1885). Deichmann (1930) misspelled the
species name as cognita. Referred to Stolus by Panning (1949). Referred
unnecessarily to a new genus Thyoneria by Deichmann (1954). We prefer
to accept the combination Stolus cognatus.

Thyone solida Deichmann, 1930. Referred to Ludwigia by Panning (1949).
Mortensen (1925) and Pawson (1963) pointed out that Ludwigia is preoc-
cupied, and is thus not available as a genus-name. Deichmann (1954) pro-
posed Euthyonacta for this species, and we accept the combination Eu-
thyonacta solida.

Thyone unisemita (Stimpson, 1851). Referred to Stereoderma by Panning
(1949).

Thyone suspecta Ludwig, 1874. Referred to Ocnus herein. Panning (1949)
had placed this species in his genus Ludwigia (see above).

Thyone surinamensis Semper, 1868. Referred to Ocnus by Panning (1949).
Named as the type of a new genus Parathyone by Deichmann (1957). We
accept the combination Ocnus surinamensis.

Thyone pervicax Theel, 1886. Referred to the preoccupied genus Lud-
wigia by Panning (1949). Referred to Thyonella by Deichmann (1954).

Thyone sabanillaensis Deichmann, 1930. In subsequent publications,
Deichmann (1941, 1954) and Panning (1949) misspelled the species-name as
sabanillensis. Referred to Thyonacta by Deichmann (1941) and by Panning
(1949), then transferred to Thyonella by Deichmann (1954). We accept the
combination Thyonella sabanillaensis.

Thyone gemmata (Pourtales, 1851). Referred to the preoccupied genus
Ludwigia by Panning (1949), and to Thyonella by Deichmann (1954).

Thyone mexicana Deichmann, 1946. Referred herein to Allothyone Pan-
ning, 1949.

394 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Genus Thyone Jaeger, 1833

Thyone Oken, 1815. (Oken’s Lehrbuch der Naturgeschichte 1815-1816
was placed on the Official Index of Rejected Works by the International
Commission on Zoological Nomenclature [Opinion 417, 1956]. As far as we
can determine, Jaeger [1833:8] was the first to validate the genus-name
Thyone in accordance with the requirements of the International Code of
Zoological Nomenclature.)

Type-species.—Holothuria fusus Miller, 1776, by original designation of
Jaeger, 1833.

Diagnosis.—Tentacles 10. Tube feet scattered on body wall, never re-
stricted to ambulacral radii. Calcareous ring with long posterior projections,
each made up of several pieces of calcite. Body wall ossicles tables with a
spire of 2 pillars. Introvert may contain rosettes only, tables only, or tables
and rosettes.

Remarks.—At the time of writing this paper, 3 species of Thyone were
known from the Western Atlantic. They are 7. inermis Heller, T. pseudo-
fusus Deichmann, and 7. pawsoni Tommasi. As T. pawsoni was described
only briefly, illustrations of the calcareous ring and of ossicles from the
introvert are given here (Fig. 2D, 2E). In this paper 2 further Thyone species
are described.

Key to Western Atlantic Species of Thyone s.s.

1. Body wall tables with oval disc, 4 perforations and thick margins

Re ee EER. Sesser Oats od EE CORI EIEIO OS 6 oo nooo 2

— Body wall tables mostly irregular in outline, perforations few to
NUMErOUS, Margin CIN A) as owe wees es Leo ore ol eee 4
2. Spires of body wall tables terminate in several short teeth ........ 3

— Spires of body wall tables terminate in a single blunt spine .......
Eee hs ee eine wes ae Thyone crassidisca, new species
3. Body wall tables with low, truncate spire. Ossicles in introvert tables

ANGETOSCELES! ee er nee Thyone pseudofusus Deichmann, 1930
— Body wall tables with high tapering spire. Ossicles in introvert ro-
SCULCSOMIV a en en oe eas Toe Thyone adinopoda, new species

4. Disc of body wall tables with 4-9 perforations. Spire of supporting
tables in tube feet abruptly tapering ... Thyone pawsoni Tommasi, 1972

— Disc of some body wall tables with 8-18 perforations. Spire of sup-
porting tables in tube feet gently tapering ... Thyone inermis Heller, 1868

Thyone inermis (Heller)

Thyone inermis Heller, 1868:78.—Deichmann, 1946:3; 1947:87, pl. 1 figs. 7—
13, pl. 2; 1954:397.—Panning, 1949:468.

VOLUME 94, NUMBER 2 395

/ ae £00)
| Wi. 00 Varo
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0) i ja
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Fig. 2. A—D, calcareous rings of some species of Thyone. Scale line for all figures = 2 mm.
A, T. adinopoda, radial and interradial pieces and single polian vesicle; B, T. crassidisca,
radial and interradial pieces and single polian vesicle; C, T. pseudofusus, two interradial pieces
and one radial; D, T. pawsoni, two interradial pieces and one radial; E, T. pawsoni, rudimen-
tary tables from introvert.

Thyone fusus: Deichmann, 1930:167, pl. 14 figs. 1-5.—Domantay, 1958:196.
Thyone deichmannae Madsen, 1941:26.

Havelockia inermis: Panning, 1949:466.

Not: Thyone fusus (Muller, 1776).

Diagnosis. —Two ventral tentacles smaller than others. Interradial pieces
of calcareous ring moderately long, extending posteriorly beyond union of
tails of radial pieces. Body wall tables with thin margins and 4 perforations,
although some have 8 or more. Spires of supporting tables in tube feet gently
tapering. Introvert with tables and rosettes.

Remarks.—Deichmann either overlooked or ignored Madsen’s new
species Thyone deichmannae, aname he applied to the W. Atlantic material
described as 7. fusus by Deichmann in 1930. If future research determines
that the Western Atlantic Thyone should be referred to a different species,
T. deichmannae Madsen is available. In the Western Atlantic, reported from
Tobago, British West Indies, and Florida (Deichmann, 1954).

Thyone pseudofusus Deichmann
Fig. 2C

Thyone pseudofusus Deichmann, 1930:168, pl. 14 tigs. 6—-9.—Clark,
1933:114.—Deichmann, 1941:107; 1946:4.—Panning, 1949:467.—Deich-
mann, 1954:395.—Domantay, 1958:197.—Tommasi, 1969:12, fig. 14.

Diagnosis.—Two ventral tentacles smaller than others. Interradial pieces

396 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

y

Fig. 3. Thyone adinopoda, ossicles. A, Tables from body wall; B, Tables from tube feet;
C, Rods, rosettes and tables from tentacles; D, Rosettes from introvert.

of calcareous ring long, extending posteriorly well beyond union of tails of
radial pieces. Body wall tables with 4 perforations, thick margins, and low
truncate spire. Introvert with tables and rosettes.

Remarks.—The type-material was from Yucatan, in 45 meters depth.

VOLUME 94, NUMBER 2 397

Deichmann (1930) also noted that the species occurred in Tobago, British
West Indies and in Florida. Tommasi reported T. pseudofusus from Sao
Paulo, Brazil.

Table 1.—Average dimensions of tables from body wall and tube feet of Thyone adinopoda.
n = 15. *, mean; SD, standard deviation; SE, standard error.

Tables from body wall

Height of table Length of table
Total length SS EEE EE Ee eee
of specimen x (um) SD SE = (um) SD SE
16 mm 82.45 11.08 2.86 98.36 8.43 2.18
22 mm 77.65 6.63 1.71 90.45 5.83 eSyll
25 mm 78.05 5.46 1.41 85.51 Se25 e§IB)
26 mm 80.25 9.69 2.50 88.59 S258) 2.48
39 mm 78.05 8.19 Petal 93.86 10.69 2.76
Supporting tables from tube feet
16 mm 62.70 8.58 DED 119.73 9.04 53)3)
22 mm 64.48 Tei 1.95 129.83 9.80 353)
25 mm 64.02 10.69 2.76 107.45 10.44 2.70
26 mm 5)3)5 10.41 2.69 130.27 12.26 3.16
39 mm 60.08" = Oaxs 1.68 117.53 8.93 2.31

Thyone pawsoni Tommasi
Fig. 2D, 2E

Thyone pawsoni Tommasi, 1972:19, figs. 12-15.

Diagnosis.—Two ventral tentacles smaller than others. Interradial pieces
of calcareous ring large, extending posteriorly well beyond union of tails of
radial pieces. Body wall tables with 4—9 perforations and thin margins. Spire
of supporting tables in tube feet abruptly tapering. Introvert with tables; no
rosettes.

Remarks.—The species was originally described from off the Gulf of Ven-
ezuela. Miller and Pawson (in preparation) describe material from off Geor-
gia and off eastern and western Florida in 24—50 meters.

Thyone adinopoda, new species
Figs. 1, 2A, 3

Diagnosis.—Two ventral tentacles smaller than others. Interradial pieces
of calcareous ring short, scarcely extending posteriorly beyond union of
tails of radial pieces. Tables in body wall with oval disc, 4 perforations,
thick margins and high tapering spires terminating in several short teeth.
Ossicles in introvert rosettes only.

398 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Etymology.—Species name derived from Greek, adinos = crowded,
pous = foot, in reference to the numerous crowded tube feet on body wall.

Material examined.—Holotype USNM E21111, total length 26 mm, U.S.
Bureau of Land Management, Sta. 13-45-VI-C(C-13), 19 July 1975,
29°30'45"N, 87°24'15"W, 63 meters. Paratypes: USNM E21112, 6 specimens,
total lengths 12, 16, 18, 22, 25, 39 mm, same locality data as holotype;
IRCZM 71:116, 2 specimens, total lengths 18, 22 mm, same locality data as
holotype. —

Description.—Body stout, more or less U-shaped, narrowing abruptly
posteriorly to form a tail. Body wall thin but moderately stiff due to nu-
merous ossicles. Tube feet numerous, apparently not completely retractile,
in double rows anteriorly and posteriorly and scattered in radii and interradii
elsewhere. Feet much more numerous in ventral interradii than in dorsal
interradii.

Body wall and tentacles dirty white to light brown in alcohol. Tentacles
10, 2 ventral smaller than others. Calcareous ring tubular (Fig. 2); radials
with very long posterior projections, each made up of numerous individual
pieces of calcite. Interradials triangular, short, scarcely extending poste-
riorly beyond union of radial tails. Interradials rounded anteriorly, radials
with anterior notch for insertion of retractor muscles. One Polian vesicle.

Body wall ossicles include tables, 60-90 um high (from bottom of half-
ring to top of spire), average 78 wm, and 72-118 um long, average 91 um.
Disc oblong, swollen medially, with smooth thick margin and 4 equal-sized
perforations. Occasionally 1—4 accessory perforations present. Spire com-
posed of 2 strong pillars joined approximately 14 the way up their length and
tapering to a blunt tip, armed with 2-7 short teeth. Opposite spire (on the
inner part of disc), projects a distinct handle or half-ring with a large central
perforation. introvert with numerous rosettes 30-100 um long, average 60
pm.
Tube feet with numerous supporting tables 50-85 um high, average 63
jm and 92-158 um long, average 120 wm. Disc elongate, strongly curved,
with 4 central perforations and usually a single perforation at each end.
Spire composed of 2 strong pillars, tapering to end in 0-6 small teeth. End
plate well developed with several concentric rings of perforations, which
increase in size from center of plate outwards.

Tentacles with numerous slender rods, rosettes and large tables. Rods
138-270 pm long, straight to slightly curved, with single perforation at each
end. Tables (average 150 um long by 80 wm wide) with flat disc, serrate
margin and numerous perforations.

Remarks.—Thyone adinopoda shares with T. crassidisca, T. pawsoni,
and 7. pseudofusus the presence of handles or half-rings on tables from the
body wall. Unlike 7. pseudofusus and T. crassidisca, T. adinopoda has
tables with a relatively thin disc, the spire is taller and gently tapering, and

VOLUME 94, NUMBER 2 399

700um

7

\ 4 °

Fig. 4. Thyone crassidisca, ossicles. A, Tables from body wall; B, Tables from tube feet;
C, Tables and rosettes from introvert; D, Table, rosettes and rods from tentacles.

400 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Average dimensions of tables from body wall and tube feet of Thyone crassidisca.
n = 15. *, mean; SD, standard deviation; SE, standard error.

Tables from body wall

Height of table Length of table

Total length S$. $<

of specimen X¥ (um) SD SE X (um) SD SE
28 mm 67.11 Told 1.84 Si=55 4.16 1.07
30 mm 70.62 Vo22 1.86 89.90 6.91 1.78
40 mm 68.42 Voll 2.01 96.48 10.75 NY
41 mm 67.11 8.33 Dal 90.79 wo1 1 35)
51 mm 78.07 11.62 3.00 94.31 8.50 2.20
60 mm 82.01 Doe) 2.56 100.45 7.65 1.98

Supporting tables from tube feet

28 mm 78.95 13.39 3.46 145.61 13.11 3.39
30 mm 83.33 18.05 4.66 144.30 16.39 4.23
40 mm 74.11 11.52 Dhl 130.71 10.52 Dee
41 mm 74.12 11.24 2.90 122.81 8.86 2S)
51 mm 91.67 9.80 2o59) 134.21 12.87 oS
60 mm 91.23 11.38 2.94 121.93 19.07 4.92

the perforations of the disc are conspicuous when viewed from above. In
T. pawsoni the handles are much less common than in T. adinopoda and
the spires of the tables are conspicuously different in both species. The
supporting tables of the tube feet in 7. adinopoda are similar to those of T.
pseudofusus. Unlike T. pseudofusus and T. crassidisca, the introvert of T.
adinopoda contains only rosettes and no tables. Like T. crassidisca, T.
adinopoda appears to have no other close relatives in the genus Thyone.

The handles or half-rings on the body wall tables of the Thyone species
discussed above are not unique to Western Atlantic species. For example,
the Australian species T. axiologa H. L. Clark, 1938, also possesses this
structure.

Thyone crassidisca, new species
Figs. 1, 2B, 4

Diagnosis.—Two ventral tentacles smaller than others. Interradial pieces
of calcareous ring moderately long, extending posteriorly well beyond union
of tail of radial pieces. Tables in body wall with oval disc, 4 perforations,
thick margins, with short spires terminating in a single blunt spine. Introvert
with tables and rosettes.

Etymology.—Species name derived from Latin, crassus = thick, discus =
plate, in reference to the thick discs of the body wall tables.

Material examined.—Holotype, USNM E21113, total length 51 mm, R/V
Gosnold Cruise 243, Station 621, 16 August 1974, 28°58.8’N, 79°58.8’W, 27

VOLUME 94, NUMBER 2 401

meters, pipe dredge, collected by D. K. Young. Paratypes: USNM E21114,
4 specimens, total lengths 28, 30, 40, 41 mm, R/V Hernan Cortez, Hourglass
Project, Station I, Field No. EJ-66-364, 4 September 1966, 26°24’N,
82°06’W, 6 meters, box dredge, sand and shell, collected by B. Presley;
USNM E21115, 1 specimen, total length 60 mm, U.S. Bureau of Land Man-
agement, Sta. HE-33-44, 18 June 1974, 28°26'30"N, 84°23’30"W, 45 meters;
IRCZM 71:117, 1 specimen, total length 59 mm, R/V Hernan Cortez, Fed-
eral Clam Project, Station 805, Field No. EJ-71-364, 23 August 1971,
25°43 .9'N, 81°42.2’W, 6.1 meters, Nantucket clam dredge, sand, 29.5°C,
collected by C. Myhree; FSBC I 2400, 1 specimen, total length 16 mm, R/V
Hernan Cortez, Hourglass Project, Station I, Field No. EJ-66-81, 9 March
1966, 26°24'N, 82°06’W, 6 meters, box dredge, mud and shell, 17.9°C, col-
lected by B. Presley.

Description.—Body fusiform, with rounded anterior and posterior ends.
Tube feet numerous, completely retractile, virtually invisible in contracted
specimens. Feet scattered over body wall, with no obvious regular arrange-
ment, although a tendency towards arrangement in double rows in radii is
apparent anteriorly and posteriorly. Body wall slightly gritty to touch. Color
whitish to light brown, with scattered light to dark brown patches of variable
size and shape, up to 1 mm in diameter. Incidence of brown patches varies;
some specimens have numerous patches, others have virtually none.

In some specimens calcareous ring, tentacles and associated structures
have been autotomized. Calcareous ring tubular, radials with long anterior
pieces and long posterior prolongations or tails. Interradial pieces also elon-
gate, extending posteriorly slightly beyond point of junction of tails. Radials
and interradials made up of a mosaic of small pieces of calcite.

Body wall ossicles include numerous tables 60-99 um high (measured
from bottom of half-ring to top of spire), average 72 wm, and 72-112 um
long, average 92 wm. Disc oval with 4 perforations, thick and strongly
knobhed on margin. Spire robust, composed of 2 pillars, truncate or greatly
tapered to a blunt tip. Inner surface of disc with distinct “‘handle’’ or half-
ring. Introvert with large tables and rosettes. Tables 92-112 um long, av-
erage 105 um, 53-60 um wide, average 56 wm and 66-92 um high, average
79 wm. Disc oval with smooth margin, 4 central perforations and several
accessory perforations at each end. Spire, composed of 2 pillars, tapers to
a single acute spine with 1—4 minute teeth at the terminus. Rosettes scat-
tered, 26-46 um long.

Tube feet contain numerous supporting tables 53-105 um high, average
82 wm and 99-165 wm long, average 133 um. Disc elongate, strongly curved,
with 4 central and 1-2 terminal perforations. Spire high, composed of 2
pillars greatly tapering to blunt or acute terminus. Endplates in tube feet
composed of several pieces.

Tentacles with rods 79-118 um long, average 90 wm with several perfo-

402 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

rations and undulate margins. Also rosettes and few tables like those found
in the introvert.

Remarks.—Thyone crassidisca appears to be most closely related to T.
pseudofusus Deichmann. They both possess robust 4-holed tables in the
body wall, the tables with a handle or half-ring on the inner side of the disc.
The species are sharply distinguished, however, on the basis of the ossicles
in the body wall and elsewhere. In the body wall tables, the spire of T.
crassidisca tends to terminate in a single point, the disc is very thick and,
when viewed from above, the disc perforations are almost completely ob-
scured by the robust spire. In 7. pseudofusus the spires are low, with a
series of (usually) 3 subequal teeth, the disc is not nearly so thick as in T.
crassidisca, and the spire is not as robust when viewed from above, so that
the disc perforations are clearly visible. The supporting tables in the tube
feet of 7. crassidisca have a spire which comes to a single blunt point, while
in T. pseudofusus the spire ends in 3—5 blunt points.

Literature Cited

Clark, H. L. 1933. A handbook of the littoral echinoderms of Porto Rico and the other West

Indian islands. Scientific Survey of Porto Rico and the Virgin Islands.—New York Acad.

Sci. 16(1):1-147, 17 pls.

. 1938. Echinoderms from Australiaa—Mem. Mus. Comp. Zool. Harvard 55(8):1-596,

63 figs., 28 pls.

Deichmann, E. 1930. The holothurians of the western part of the Atlantic Ocean.—Bull. Mus.

Comp. Zool. 71(3):41-226, 24 pls.

. 1941. The Holothuriodea collected by the Velero III during the years 1932-1938. Part

1, Dendrochirota.—Allan Hancock Pacific Exped. 3(83):61—-194, 21 pls.

. 1946. A new species of Thyone s.s. from the Gulf of Mexico.—Occas. Pap. Mar.

Biol. Lab., Louisiana State Univ., Baton Rouge, La. No. 4:1-4, 1 fig.

. 1947. The fusus-like Thyone from the West Indian waters.—Proc. New England Zool.

Club 24:83-90, 2 pls.

. 1954. The holothurians of the Gulf of Mexico. In Gulf of Mexico, its origin, waters

and marine life.-—U.S. Fish Wildl. Serv. Fish. Bull. 89:381—410, 3 figs.

———. 1957. The littoral holothurians of the Bahama Islands.—American Mus. Novit.

1821:1-20.

. 1963. Shaliow-water holothurians known from Caribbean waters.—Stud. Fauna Cu-

racao 14(63):100-118.

Domantay, J. S. 1958. Some holothurians from Florida in the collection of the Allan Hancock
Foundation.—Agra University J. Res. Sci. 7(2):181-202.

Heller, C. 1868. Die Zoophyten und Echinodermen des Adriatischen Meeres.—K.K. Zool.-
Bot. Ges. Vienna: 1-88, 3 pls.

Jaeger, G. F. 1833. De Holothuriis. Turin. Pp. 1-42, 3 pls.

Lampert, K. 1885. Die Seewalzen. Weisbaden- Pp. 1-312, 1 pl.

Lesueur, C. A. 1824. Description of several new species of Holothuria.—Jour. Acad. Nat.
Sci. Philad. 4:155-163.

Ludwig, H. 1874. Beitrage zur Kenntnis der Holothurien.—Arb. Zool. Zoot. Inst. Wurzburg
3:77-120, 2 pls.

VOLUME 94, NUMBER 2 403

. 1887. Die von G. Chierchia auf der Fahrt der Kgl. Corvette Vettor Pisani gesammelten

Holothurien.—Zool. Jahrb. 2:1—36, 2 pls.

Madsen, F. J. 1941. On Thyone wahrbergi n.sp., a new holothurian from the Skagerrak, with
remarks on T. fusus (O.F.M.) and other related species—Goteborgs VetenskSamh.
Handl. (6) 1B 1:1-31, 17 figs.

Mortensen, T. 1925. Echinoderms of New Zealand and the Auckland and Campbell Islands.
Part IV. Holothurioidea.—Vidensk. Medd. Dansk Naturh. Foren. 79:322-386.

Miiller, O. F. 1776. Prodromus Zoologiae Danicae seu animalium Daniae et Norvegiae Indi-
genarum Characters, Nanina et Synonyma. Imprimis Popularium Hauniae. [Holothuri-
ans p. 231-232.]

Panning, A. 1949. Versuch einer Neuordnung der Familie Cucumariidae. (Holothu-
roidea, Dendrochirota).—Zool. Jahrb. 78(4):404—470, 62 figs.

Pawson, D. L. 1963. The holothurian fauna of Cook Strait, New Zealand.—Zool. Publ. Vic-
toria Univ. Wellington 36:1-38, 7 pls.

Pourtales, L. F. De. 1851. On the Holothuriae of the Atlantic coast of the United States.—
Proc. Am. Assoc. Adv. Sci. :8—16.

Semper, C. 1868. Reisen in Archipel der Philippinen, Theil II. Wiss. Res. Bd. 1, Ho-
lothurien. Pp. 1-228, 40 pls.

Sluiter, C. P. 1910. Westindischen Holothurien.—Zool. Jahrb. f. Anat. und Syst. 11(2):331-
341.

Stimpson, W. 1851. Pp. 7-8 in Proceedings of January 15, 1851.—Proc. Boston Soc. Nat.
Hist. IV.

Theel, H. 1886. Report on the Holothuriodea of the Blake Expeditions 1887—1880.—Bull.
Mus. Comp. Zool. 13(1):1—21, 2 pls.

Tommasi, L. R. 1969. Lista dos Holothurioidea recentes do Brasil. Contrcoes.—Inst. Ocean-

ogr. Univ. S. Paulo, ser. Ocean. Biol. 15:1—29, 27 figs.

. 1972. Equinodermes do Regiao Entre O Amapa (Brasil) E.A. Florida (E.U.A.).—

Bolm. Inst. Oceanogr. S. Paulo 21:15-—67, 27 figs.

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(DLP) Department of Invertebrate Zoology, National Museum of Natural
History, Washington, D.C. 20560; (JEM) Harbor Branch Foundation, Inc.,
Route 1, Box 196, Fort Pierce, Florida 33450.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 404-412

TWO LITTLE-KNOWN SPECIES OF
ELEUTHERODACTYLUS (AMPHIBIA:
LEPTODACTYLIDAE) FROM THE SIERRA
DE LA MACARENA, COLOMBIA

William F. Pyburn and John D. Lynch

Abstract.—Two sympatric species of Eleutherodactylus occur in the
cloud forest of La Macarena, Colombia. One is an undescribed species,
herein named Eleutherodactylus savagei. This species resembles E. fenes-
tratus, from which it differs in skin texture, in having a heel tubercle, and
in throat pattern.

The other Macarena species is Eleutherodactylus frater. The species E.
savagei and E. frater are morphologically similar and have similar breeding
calls, but appear to be reproductively isolated from each other by differ-
ences in habitat, period of activity, and possibly by subtle differences in
call.

A field party from the University of Texas at Arlington collected speci-
mens and obtained voice recordings of two species of Eleutherodactylus in
the cloud forest of the Macarena Mountains, Departamento del Meta, Co-
lombia, in March 1971. The calls were recorded with a Uher 4000 Report
L tape recorder and sound spectrograms were made from the tapes in the
laboratory with a Kay Elemetrics Corporation ‘‘Sona-Graph.”’

One of the species is Eleutherodactylus frater, a small arboreal form
whose relationships have been in doubt. The other species is unnamed. For
it, we propose the name:

Eleutherodactylus savagei, new species
Gigs)

Holotype.—UTA 3535, an adult male taken on the Sierra de la Macarena,
ca. 35 km WSW Vistahermosa, Depto. Meta, Colombia, 1097-1128 m, on
20 March 1971 by W. F. Pyburn.

Paratypes.—UTA 3525-29, 3536-38, 5608-18; USNM 218054-62;
AMNH 107557-65; CM 76154-62; KU 187001-09; all topotypes collected
17-20 March by W. F. Pyburn, J. R. Glidewell, and J. Taulman.

Diagnosis.—1) Skin of dorsum finely shagreened becoming more coarse
posteriorly, that of venter smooth; no dorsolateral folds; 2) tympanum prom-
inent, its length about '4 eye length; 3) snout subacuminate in dorsal view,

VOLUME 94, NUMBER 2 405

Fig. 1. Hands of (A) Eleutherodactylus savagei n. sp., UTA 5616, and (B) E. frater, UTA
3531. Lines equal 2 mm.

acutely rounded in lateral profile; snout long, E-N < eye length; 4) IOD
broader than upper eyelid, flat; 5) vomerine odontophores triangular in out-
line; 6) males with vocal slits and white, non-spinous nuptial pads; 7) length
of first and second fingers nearly equal; large pads on fingers IIIJ-IV; 8)
fingers with slight lateral keels; 9) only antebranchial tubercle evident; 10)
small tubercles on heel; no outer tarsal tubercles; inner edge of tarsus bear-
ing ridge or series of tubercles; 11) two metatarsal tubercles, inner elongate,
4 times size of outer; supernumerary plantar tubercles at bases of toes; 12)
toes bearing keels proximally; basally webbed (web not encroaching basal
subarticular tubercles); 13) tan to brown with pattern vague except for black
scapular spots and canthal-supratympanic stripe; throat peppered with
brown, pigment extending onto chest as faint reticulation; groin and anterior
and posterior surfaces of thighs uniform brown; shank bars oblique, nar-
rower than interspaces; 14) adults moderate-sized, males 17.7—22.6 mm SVL
Ges— 9.02 Ne.) females. 3 0.0—34 J.mm.(x =) 52-6, No—, 17).

Most similar to E. fenestratus (Steindachner) but differing in skin texture
(skin of dorsum finely shagreened vs with an interspersion of large warts),
in having a small heel tubercle (none in E. fenestratus), and in having only
faint marbling on the throat.

Description.—Head narrower than body, wider than long; HW in males

406 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Color pattern of Eleutherodactylus savagei n. sp., UTA 3527. Line equals 10 mm.

33.4-39.4 (x = 37.2, N = 5) percent SVL, in females 37.7—44.2 (x = 40.2,
N = 18) percent; snout subacuminate in dorsal view, acutely rounded in
lateral profile; snout long, E—N in males 73.5—75.8 ( = 74.4, N = 4) per-
cent eye length, in females 79.5-95.0 (x = 86.9, N = 20) percent; nostrils
weakly protuberant, directed laterally; canthus rostralis sharp, nearly
Straight (sinuous); loreal region flat (or very weakly concave), sloping
abruptly to lip; some flaring of jaws posteriorly; interorbital region flat, no

VOLUME 94, NUMBER 2 407

cranial crests; upper eyelid width in males 83.0-100.0 (x = 89.9, N = 4)
percent IOD, in females 65.8—109.4 (< = 80.1, N = 8) percent; supratym-
panic fold thick, obscuring upper and posterior edges of tympanum which
is higher than long, length of tympanum in males 29.3—33.3 (¢ = 32.1, N =
4) percent eye length, in females 27.5-35.7 (x = 32.3, N = 20) percent;
postrical tubercles not prominent; choanae round, moderately large, not
concealed by palatal shelf of maxillary arch; vomerine odontophores slightly
smaller than a choana, triangular in outline, separated on midline by a dis-
tance equal to odontophore width, median and posterior to choanae, each
bearing 4—5 teeth; tongue slightly longer than wide, not notched posteriorly,
posterior 4 not adherent to floor of mouth; males with vocal slits and a
subgular vocal sac.

Skin of dorsum finely shagreened anteriorly, becoming more coarse pos-
teriorly and very coarse with scattered tubercles on flanks; 1-2 small tu-
bercles on upper eyelid; some evidence of a dorsolateral fold in scapular
region but fading out in region of sacrum; large areolations below and pos-
terolateral to vent, on posteroventral surfaces of thighs; no anal sheath or
enlarged para-anal warts; skin of throat and venter smooth but with some
suggestion of areolations encroaching from flanks; discoidal folds ill-defined;
no ulnar tubercles except antebrachial; palmar tubercle bifid, much larger
than oval thenar tubercle; supernumerary palmar tubercles relatively large
(Fig. 1); subarticular tubercles subconical, round or slightly longer than
wide; fingers bearing faint lateral keels; finger tips expanded forming pads
(smallest on thumb), bearing broad discs on ventral surfaces; pads of IIJ—
IV largest (as large as tympanum) truncate, those of I-II round (Fig. 1);
-males bearing non-spinous nuptial pad; length of first and second fingers
nearly equal.

One to two low round tubercles on heel, none on outer edge of tarsus;
inner edge of tarsus bearing a long tubercle (sometimes one long and a short
proximal one); inner metatarsal tubercle three times as long as wide; outer
metatarsal tubercle round, 4 size of inner; supernumerary plantar tubercles
4 (rarely 5), at bases of toes (some hint of others on rest of sole); basal toe
webbing (not encompassing basal subarticular tubercles); lateral keels evi-
dent low on toes but indistinct distally; toes bearing pads and broad discs
(smaller than those of outer fingers); heels of flexed hind legs overlap slight-
ly; heel of adpressed hind leg reaches anterior edge of eye; shank of males
51.5—58.0 (« = 54.5, N = 5) percent SVL, of females 50.0—58.0 (* = 53.9,
N = 20) percent.

Dorsum tan to brown with black scapular spots and nearly as dark can-
thal-supratympanic stripes; light brown sacral chevron, suprainguinal
marks, interorbital bar, and flank bars (Fig. 2); anal triangle brown outlined
with cream; 3—4 oblique bars on shank, less than to equal interspace width;
a common variant has cream dorsolateral stripes along anterior 37/s body

408 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

0.2 0.4 0.6 0.8 1.0 1.2
seconds

Fig. 3. Sound spectrogram of one complete call of Eleutherodactylus savagei (narrow band
filter). Recorded 20 March 1971 at type locality (air temperature, 17.3°C).

(Fig. 2); throat lightly peppered with brown, sometimes extending onto chest
as a reticulation; peppering on undersides of legs; groin and anterior and
posterior surfaces of thighs uniform brown.

Measurements of holotype in mm.—SVL 21.2; shank 11.5; head width
7.8; head length 8.0; upper eyelid width 2.4; IOD 2.4; tympanum length 1.1;
eye length 3.4; E—N 2.5.

Voice.—In the Macarena Mountains, Eleutherodactylus savagei began
calling at dusk and choruses continued until well after midnight. The males
called from low perches on leaves up to 50 cm above the ground. The call
(Fig. 3) was a series of short closely spaced notes forming a dominant fre-
quency band at 3000 Hz. Nineteen calls of 7 frogs recorded 20 March 1971
(air temperature, 17.3°C) had an average of 15 (range, 7-34) notes per call
and a note repetition rate of 11.3 (10.2—12.2) notes per s. The duration of
the call was 1.3 s (0.6—3.2) and the note duration was 0.05 s (0.04—0.09).

Etymology.—The new species is named for Jay M. Savage of the Uni-
versity of Southern California, in recognition of his contributions to under-
standing the biology of Neotropical Eleutherodactylus.

Distribution.—In addition to the type-series, we have seen the following
(also from Depto. Meta, Colombia): Buenavista, 10 km NW Villavicencio,
UTA 2723, 2725-26, 2741, 2743, 2748; Serrania de la Macarena, FMNH
81349-60, 81364.

Remarks.—Cochran and Goin (1970) confused this frog among what they
called E. terraebolivaris Rivero (which occurs only in Venezuela). Lynch
(1975) noted the presence of this frog but mistakenly termed it a member of
the unistrigatus group. Eleutherodactylus savagei does not appear closely

VOLUME 94, NUMBER 2 409

allied to the short-legged fitzingeri group of frogs of the Amazon Basin (E.
lanthanites and E. vilarsi). It seems to be the northwestern replacement for
E. fenestratus (Steindachner), which is replaced in northeastern South
America by E. gutturalis Hoogmoed, Lynch, and Lescure.

Eleutherodactylus frater (Werner)

Hylodes frater Werner, 1899, Verh. zool.-bot. Gessell. Wien, 49:479 (type-
locality, Peperital, near Villavicencio, Depto. Meta, Colombia, 1000 m).

Eleutherodactylus frater: Cochran and Goin, 1970, Bull. U.S. Nat. Mus.
(288):433-34, pl. 54.

Diagnosis.—1) Skin of dorsum finely shagreened with scattered warts,
that of venter coarsely areolate; no dorsolateral folds; 2) tympanum prom-
inent, its length 4-7/5; eye length; 3) snout subacuminate in dorsal view,
round in lateral profile; canthus rostralis obtuse; 4) upper eyelid as wide as
IOD; no cranial crests; tubercles on upper eyelid; 5) vomerine odontophores
small, low; 6) males with vocal slits, subgular vocal sac; males bearing non-
spinous nuptial pads on thumbs; 7) first finger shorter than second; large
pads on outer fingers; 8) fingers with narrow lateral keels; 9) 2-3 conical
ulnar tubercles present; 10) small tubercles on heel; inner edge of tarsus
bearing short ridge, outer edge with 2-3 conical tubercles; 11) two metatar-
sal tubercles, inner oval, 8 times size round outer; supernumerary plantar
tubercles present; 12) toes bearing lateral keels, enlarged pads; 13) pale
brown with brown markings (reverse parentheses, suprainguinal blotches,
narrow limb bars, supratympanic stripe and labial bars); no canthal stripe;
venter finely peppered with brown (throat darker); concealed surfaces of
thighs uniform brown; 14) adults small, males 17.2—21.6 mm SVL (% = 20.1,
N = 7), one adult female 24.9 mm SVL.

Eleutherodactylus frater is most similar to E. incomptus Lynch and Duell-
man, E. marmoratus (Boulenger), and E. martiae Lynch, but differs from
each in the arrangement of tarsal tubercles. It also differs from the smaller
E. martiae in having vocal sacs in males and prominent tympani (concealed
in E. martiae). Eleutherodactylus incomptus is probably the closest relative
of E. frater but is slightly smaller (males 15.6-18.8 mm, females 23.7—25.9
mm SVL) and has the vomerine odontophores evident only in large females.
Eleutherodactylus marmoratus is considerably larger (its males are larger
than our only female E. frater).

Description.—Head as wide as body, wider than long; HW 35.4—38.4 (x =
36.8, N = 7) percent SVL; snout subacuminate in dorsal view, round in
lateral profile; nostrils weakly protuberant, directed dorsolaterally; snout
short, E—N 69.2-87.3 (x = 80.1, N = 7) percent eye length; canthus ros-
tralis weakly defined, rounded; loreal region weakly concave, sloping to
lips, lips not flared; upper eyelid width 93.3-113.6 (x = 101.6, N = 7) per-

410 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

cent IOD; upper eyelid bearing 2—4 conical tubercles; no cranial crests;
supratympanic fold concealing upper edge of tympanic annulus; tympanum
prominent, round, its length 26.9-38.5 (<« = 33.4, N = 7) percent eye length,
separated from eye by slightly more than its diameter; postrictal tubercles
subconical; choanae relatively small, round, not concealed by palatal shelf
of maxillary arch; vomerine odontophores small, each 4%4—% size of a
choana, narrowly separated (by distance equal 4 width of an odontophore),
bearing 2-3 teeth; tongue about as long as wide, posterior border feebly
indented, posterior 2/s not adherent to floor of mouth; males with relatively
long vocal slits and subgular, external vocal sac.

Skin of dorsum finely shagreened, overlain with larger granulations on
lower back and with larger isolated tubercles on upper eyelid, between eyes,
and in scapular region; flanks becoming coarsely areolate ventrally, skin of
venter areolate, that of throat smooth; no dorsolateral folds or anal sheath;
discoidal folds prominent; 2-3 ulnar tubercles; palmar tubercle bifid, much
larger than oval thenar tubercle (Fig. 1); supernumerary palmar tubercles
smaller than more pungent, round subarticular tubercles; fingers bearing
slight lateral keel, most obvious along outside of palm and finger IV; pads
truncate, smallest on I, that on II intermediate, largest on III and IV (more
than 2 times width of digit below pad); pad of III as large as tympanum;
discs broader than long, not indented, first finger shorter than second (Fig.
1); males with white, non-spinous nuptial pad on thumb.

Small tubercle on heel, 2-3 on outer edge of tarsus, inner edge of tarsus
bearing tarsal ridge (distal 2) and a tubercle proximal to ridge; inner meta-
tarsal tubercle twice as long as wide, outer minute (less than 1% size of
inner), round; supernumerary plantar tubercles small, pungent, toward dis-
tal end of sole; subarticular tubercles round, weakly pungent; toes bearing
lateral keel (more fringe-like than that on fingers) and pads (about same size
as on fingers but seem smaller because of toe fringe); heels of flexed hind
legs barely overlap; shank 46.8—50.0 (x = 48.9, N = 7) percent SVL.

Pale brown with slightly darker brown marbling over dorsum; still darker
reverse parentheses in scapular region, slanted bars on upper flanks, and
suprainguinal blotches; anal triangle dark brown; limb bars narrower than
interspaces, oblique on shanks; no canthal stripe; supratympanic stripe dark
brown, as are labial bars; diffuse brown area behind shoulder; ventral sur-
faces finely dusted with brown (most dense on throat) but appearing cream
without magnification; groin peppered with brown, anterior and posterior
surfaces of thighs uniform brown.

UTA 3531 has a black dorsolateral stripe continuing anteriad as a canthal
stripe. There are 3 shorter brown stripes down the center of the back. Some
specimens also have very pale snouts and, if so, the pale area defines an
otherwise indistinct interorbital bar.

Voice.—During our 7-day camp in the cloud forest, choruses of E. frater

VOLUME 94, NUMBER 2 411

seconds

Fig. 4. Sound spectrogram of incomplete call (last 19 notes) of Eleutherodactylus frater
(narrow band filter). Recorded 21 March 1971, Sierra de la Macarena, Colombia (air temper-
ature, 19.7°C).

began each day at about 1500 h and continued through the evening until
dark. Most individuals became quiet at dusk but a few frogs continued to
call until about | hour after dark.

The call (Fig. 4) was a series of well-separated, short notes with a dominant
frequency of about 2900 Hz. Thirteen calls of 4 individuals recorded 21
March 1971 (air temperature, 19.7°C) had a mean of 17.7 (11-32) notes per
call and a note repetition rate of 16.7 (14.8-18.1) notes per s. The duration
of the call was 1.1 s (0.7-2.2) and the note duration was 0.03 s (0.02—0.06).

Remarks.—Cochran and Goin (1970) confounded E. frater with the cho-
coan E. taeniatus (Boulenger). Lynch (1974) used the name E. frater for
the chocoan species but subsequently redescribed E. taeniatus (Lynch,
1980). The eastern face of the Cordillera Oriental in Colombia is poorly
known herpetologically but we anticipate E. frater and E. incomptus to
replace one another somewhere between the Pasto-Mocoa road and the
Bogota-Villavicencio road.

Discussion

Eleutherodactylus frater and E. savagei are similar enough in their mor-
phology and breeding calls to make one suspect that cross matings could
occur. Eleutherodactylus savagei begins calling at dusk, about 1 h before
dark, so there is a 2 h interval in which the two species call at the same
time. However, E. frater is almost entirely diurnal and E. savagei is essen-
tially nocturnal, there being relatively few individuals of either species that
call in the interval of temporal overlap. Furthermore, E. frater is arboreal
whereas E. savagei males call from perches near the ground.

412 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Both species produce a varying number of short notes with about the
same dominant frequency, but calls of the two species can easily be distin-
guished by the human ear. The sound spectrograms show that the note
duration of E. savagei averages nearly twice that of E. frater and that the
notes of E. savagei are repeated at a slower rate than the notes of E. frater.
We would expect these differences in habitat, breeding call, and time of
vocalization to function as premating isolation mechanisms, making matings
between the two species unlikely.

Abbreviations

Institutional abbreviations: AMNH, American Museum of Natural His-
tory, New York; FMNH, Field Museum of Natural History, Chicago; KU,
University of Kansas Museum of Natural History, Lawrence; USNM, Na-
tional Museum of Natural History, Washington, D.C.; UTA, University of
Texas at Arlington, Arlington, Texas; CM, Carnegie Museum, Pittsburgh,
Pennsylvania.

Acknowledgments

We thank Hymen Marx of the Field Museum of Natural History for the
loan of specimens, and Wanda C. Pyburn, Jerry Glidewell, and James Taul-
man for assistance in the field. Travel by the senior author was made pos-
sible by a grant from the National Science Foundation in support of U.S.
participation in the International Biological Program. Officials of the Insti-
tuto Nacional de Recursos Naturales (INDERENA) gave permission to col-
lect specimens in the Sierra de la Macarena. Representative speimens and
color transparencies of habitats were deposited in the collections of IN-
DERENA.

Literature Cited

Cochran, D. M., and Goin, C.J. 1970. Frogs of Colombia.—U.S. Nat. Mus. Bull. (288):i—xii +
1-655, 68 pls.

Lynch, J. D. 1974. "New species of frogs (Leptodactylidae: Eleutherodactylus) from the Am-
azonian lowlands of Ecuador.—Occ. Pap. Mus. Nat. Hist. Univ. Kansas (31):1-22.

. 1975. The identity of the frog Eleutherodactylus conspicillatus (Gunther), with de-
scriptions of two related species from northwestern South America (Amphibia, Lepto-
dactylidae).—Contrib. Sci., Nat. Hist. Mus. Los Angeles Co. (272):1-19.

. 1980. Systematic status and distribution of some poorly known frogs of the genus Eleu-
therodactylus from the chocoan lowlands of South America.—Herpetologica 36: 175-189.

(WFP) Department of Biology, The University of Texas at Arlington,
Texas 76019; (JDL) School of Life Science, The University of Nebraska,
Lincoln, Nebraska 68588.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 413-420

THE SYSTEMATIC STATUS OF CENTRAL
AMERICAN FROGS CONFUSED WITH
ELEUTHERODACTYLUS CRUENTUS

Jay M. Savage

Abstract.—Eleutherodactylus cerasinus, E. cruentus, and E. ridens, three
superficially similar species from lower Central America, are re-diagnosed.
Direct comparisons of type material show that E. peraltae Barbour, 1928,
and E. tiptoni Lynch, 1964, are synonyms of E. cerasinus; Syrrhophus
lutosus Barbour and Dunn, 1921, E. dubitus Taylor, 1952, and E. marshae
Lynch, 1964 are synonyms of E. cruentus; and Syrrhopus molinoi Barbour,
1928 is a synonym of E. ridens.

A series of three superficially similar species of Eleutherodactylus from
lower Central America have caused confusion for all workers on the genus.
The forms share the following features: finger II longer than I; no toe web-
bing; large finger and toe disks; disk pads round, never triangular; disk
covers round, palmate or truncate, never pointed; and venter coarsely are-
olate. The earliest name applied to these frogs is Hylodes cruentus W.
Peters, 1873. Since that time eleven different names, eight proposed as new
taxa, have been applied to these forms.

In the course of studies on the genus in Middle America, I have had the
opportunity to see extensive new living and preserved material of the prob-
lematic populations and have examined all extant types of names involved
in the problem. The following sections summarize my findings as a basis for
a more extensive treatment of all Middle American species in the genus in
a subsequent paper. The clarification of the taxonomy and synonymy for
these forms is required at this time so that other workers may use the correct
names in their systematic and ecological treatments.

Eleutherodactylus cruentus (W. Peters)

1873 Hylodes cruentus W. Peters, Mon. Akad. Wiss. Berlin: 609.

1921 Syrrhophus lutosus Barbour and Dunn, Proc. Biol. Soc. Wash. 34:158
(Holotype: MCZ 8023; Costa Rica: Cartago: Canton de Cartago: Na-
varro, 1100 m).

1931 Eleutherodactylus ventrimarmoratus: Dunn, Occ. Paps. Boston Soc.
Nat. Hist. 5:411.

Eleutherodactylus lutosus lutosus: Dunn (by implication), Occ. Paps.
Boston Soc. Nat. Hist. 5:411.

414 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1933 Eleutherodactylus latidiscus: Dunn, Occ. Paps. Boston Soc. Nat.
Hist. 8:68-69.

1952 Eleutherodactylus dubitus Taylor, Univ. Kansas Sci. Bull. 35(5):707,
Fig. 22 (Holotype: KU 24942; Costa Rica: Alajuela: Canton de Ala-
juela: SE slope Volcan Poas: Isla Bonita, 1476 m).

1964 Eleutherodactylus marshae Lynch, J. Ohio Herp. Soc. 4(3):67, Fig.
3 (Holotype: BYU 18819; Panama: Bocas del Toro: Rio Changena, 35
km W Almirante).

1965 Eleutherodactylus cruentus: Savage, Bull. South. Calif. Acad. Sci.
64(2):109.

Holotype.—Berlin 7811; Panama: Provincia de Chiriqui.

The confusion surrounding proper application of the name cruentus stems
from several sources. Two of these are biological. The first is the result of
extensive sexual dimorphism in which the males are much smaller (maxi-
mum size 28 mm) than the females (to 42 mm), and are also much more
tuberculate and tend to have a truncate snout often with a slightly fleshy
pointed tip (females are smoother and usually with a subovoid snout in
dorsal outline). In addition, the tympanum in males is completely hidden by
skin, whereas in females it is indistinct but definitely visible. The second
involves the extremely high variability in coloration, ranging from markedly
patterned to nearly uniform tan to black in dorsal color, often with light
areas on back, flanks, and thighs green, bright yellow, orange or red; groin
usually with one to several large yellow to orange spots. Nevertheless, all
individuals assigned to this species agree in having a distinct, enlarged,
pointed or flap-like supraocular tubercle; a well developed, distinct, pointed
heel tubercle; greatly expanded, truncate, emarginate finger disks; and the
subarticular tubercles raised but not pointed.

The holotype of this species is an adult female, 39.5 mm in length, col-
lected by Ribbe somewhere in the Chiriqui region of western Panama. It
agrees in every respect (see key couplets | and 2 below) with examples
now known from Costa Rica and Panama, although the extensive suffusion
of the dorsum with red (the source of the species name) described by W.
Peters (1873) is an unusual variant. Recently collected female examples from
above Boquete, Chiriqui Province, Panama, are often colored in a similar
fashion.

When the unusual coloration of the type of cruwentus and the marked
sexual dimorphism of the species are taken into account, it is not surprising
that Barbour and Dunn (1921) described Syrrhophus lutosus based upon a
juvenile male, 13 mm in length. This example agrees in all ways with male
cruentus. Difficulty arose later, however, when Dunn (1931) allied lutosus
with the nominal species Syrrhopus [sic] molinoi Barbour (1928), by reduc-
ing the latter to subspecies status under /utosus. As we shall see later,
molinoi belongs with a very different species population.

VOLUME 94, NUMBER 2 415

While confused about the relationships of *‘lutosus’’ and molinoi Dunn
(1931, 1933) was well aware that molinoi differed from the species here
called cruentus. He referred material of the latter from central and western
Panama, first to Eleutherodactylus ventrimarmoratus (an Amazonian form)
and then to E. latidiscus (a western South American species). At the same
time he recognized a relative of cruentus, Eleutherodactylus cerasinus
(Cope, 1875), as occurring on Barro Colorado Island, Panama. Eleuthero-
dactylus cerasinus is distinctive in having the groin, posterior thigh, and
calf suffused with red pigment in life, a condition never found in cruentus.

Sometime between 1931 and 1951, Dunn (in Zetek and Wetmore, 1951)
apparently became convinced that cruentus was an older name for cerasi-
nus. He thus applied the former to cerasinus, while using latidiscus for
cruentus. As aresult almost all examples of cerasinus at most United States
museums were re-labeled cruentus by Dunn. One suspects the change came
about because Dunn wanted to apply the old name cruentus to some Panama
form. Although he had examined the type of cruentus at Berlin in 1928, his
notes may have been inadequate and his memory somewhat diminished by
the 23 year gap. Very likely the red on the thighs of cerasinus and the name
cruentus (= bloody) influenced him. As we shall see below, the two names
apply to distinctive species.

Taylor (1952) in his classic work on Costa Rica recognized Jutosus and
cerasinus as distinct. In addition he described as new Eleutherodactylus
dubitus, based on a large and variable series including males and females,
but with a female holotype, 38.5 mm in length. Taylor’s description covers
a fair proportion of the variation in the species, but direct comparison of
his type and the type of cruentus leaves no doubt as to their conspecificity.

Eleutherodactylus marshae Lynch, 1964 is a poorly fixed (the type and
only known example appears to have died sometime well before preserva-
tion) adult male, 28 mm in length. The distinct black bars and bright yellow
groin spots confirm the identification with cruentus made on the basis of
direct comparison to Costa Rica and Panama material. The smooth venter
and apparent tympanum mentioned in the original description are artifacts
of death and desiccation prior to preservation. The type of marshae further
agrees with adult male cruentus in having nuptial thumb pads and lacking
vocal slits.

Eleutherodactylus cruentus differs from other Central American members
of the genus with which it might be confused (exclusive of the two forms
discussed in detail later in this paper) in having a well developed pointed to
flap-like heel tubercle or calcar (absent in E. altae of Costa Rica and E.
pardalis of Costa Rica and Panama); in having well developed accessory
palmar tubercles and a series of 2—4 outer tarsal tubercles (absent in E.
caryophyllaceus and E. melanostictus of Costa Rica and western Panama);
and in having a distinct well developed pointed to flap-like supraocular tu-

416 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

bercle (absent in taeniatus of Panama, Colombia, and northwestern Ecua-
dor).

The known geographic range of this species may be stated as follows:
primarily humid forests in the premontane zone on both slopes of Costa
Rica and western Panama; scattered lowland records in eastern and south-
western Costa Rica into central Panama (40-1550 m).

Referral of eastern Panama and Colombia material to this species (Lynch
1979) is subject to question and requires a detailed study and comparison
of samples before their status can be resolved.

Eleutherodactylus cerasinus (Cope)

1875 Hylodes cerasinus Cope, J. Acad. Nat. Sci. Phila. 2(8):112, pl. 26,
Fig. 4.

1921 Eleutherodactylus cerasinus: Barbour and Dunn, Proc. Biol. Soc.
Wash. 34:161; 1952, Taylor, Univ. Kansas Sci. Bull. 35(5):703, Fig.
21. |

1928 Eleutherodactylus peraltae Barbour, Proc. N. Engl. Zool. Club 10:30,
pl. 4, Fig. 1 (Holotype: MCZ 13601; Costa Rica: Cartago: Canton de
Turrialba: Peralta, 386 m).

1964 Eleutherodactylus tiptoni Lynch, J. Ohio Herp. Soc. 4(3):65, Fig.
2 (Holotype: BYU 19146; Panama: Bocas del Toro: 35 km W Almi-
rante).

1951 Eleutherodactylus cruentus: Dunn in Zetek and Wetmore, Ann. Rept.
Smithson Inst. 1950:140.

Holotype. —USNM 32572: Costa Rica: Provincia de Limon: Canton de
Talamanca: Cerro Ujam, 1524-2134 m.

This species was described by Cope (1875) based on material from the
expedition of William More Gabb to Cerro Ujam in southeastern Costa Rica.
The holotype, an adult female 33 mm in length, is still extant but is in very
poor condition. The frog clearly was not collected at the elevation indicated
in the original description (Savage, 1970). This specimen has all the distinc-
tive features of cerasinus: a well developed heel tubercle; no enlarged su-
praocular or superciliary tubercles; some subarticular tubercles under toes
III-IV pungent to pointed; tympanum clear, distinct; no discrete light and
dark markings on groin. The presence of red in the groin, on the posterior
surface of the thigh, and on the inner face of the calf as described by Cope
(1875) confirms the facts of morphology as supported by the original figure.
The species was independently described by Barbour (1928) as Eleuthero-
dactylus peraltae based on a juvenile 26 mm in length. This individual agrees
in all diagnostic features with the type of cerasinus and material from lo-
calities near Peralta on the Atlantic slope of Costa Rica.

VOLUME 94, NUMBER 2 417

The type and only known specimen of E. tiptoni is a young female 24 mm
in length, that appears to have died and become desiccated before preser-
vation. Its distinctive features include: a well developed heel tubercle, a
series of subequal superciliary tubercles, outer finger disks expanded but
not emarginate and some subarticular tubercles pungent or pointed. In these
features the type of tiptoni agrees with E. cerasinus as defined in this paper
and approaches closely examples referred to the latter form from the AI-
mirante area of northwestern Panama.

Eleutherodactylus cerasinus differs from E. cruentus in the many features
given in the key below (couplet 2). The types of cerasinus, peraltae and
tiptoni conform closely with this character suite. Dunn (in Zetek and Wet-
more, 1951) used the name cruentus for an Eleutherodactylus population
from Barro Colorado Island, Panama. A re-examination of material in many
museum collections from the island and other areas in Costa Rica and Pan-
ama, identified by Dunn as cruentus, shows that he consistently misapplied
the latter name to examples of E. cerasinus. Although Myers and Rand
(1969) continued to use cruentus for Barro Colorado Island examples with
some question, the species is not known to occur on the island and all
references to its occurrence there are based on cerasinus.

Among Central American Eleutherodactylus that might be mistaken ini-
tially for E. cerasinus, each (beside the two forms reviewed in greater detail
in this paper) may be readily separated from it as follows (features for
cerasinus in parentheses): E. altae and E. pardalis lack a well developed
heel tubercle and have large discrete light groin and thigh spots (tubercle
present, no discrete light spots); E. caryophyllaceus has an enlarged su-
perciliary tubercle (no enlarged superciliary tubercle, although a row of
equal sized small superciliary tubercles usually present); E. melanostictus
has a well developed supraocular tubercle (absent); and E. taeniatus has
distinct narrow finger fringes, lacks superciliary tubercles, lacks red in the
groin and on legs in life, and adult males have nuptial thumb pads (no distinct
finger fringes, a series of small superciliary tubercles, red present in the
groin and on legs in life, no nuptial thumb pads).

The known geographic range for this species is: the Atlantic lowlands and
premontane slopes of Nicaragua, Costa Rica, and Panama, onto both ver-
sants in western and central Panama (40-1300 m).

Eleutherodactylus ridens (Cope)

1867 Phyllobates ridens Cope, Proc. Acad. Nat. Sci. Phila. 18:131.

1885 Hypodictyon ridens: Cope, Proc. Amer. Philos. Soc. 22:383.

1888 Syrrhophus ridens: Boulenger, Proc. Zool. Soc. London: 206.

1900 Syrrhophus ridens: Gunther, Biol. Centr. Amer.: 216.

1928 Syrrhopus molinoi Barbour, Proc. N. Engl. Zool. Club 10:28 pl. 2,

418 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2 (Holotype: MCZ 13051: Panama: Canal Zone: Barro Colorado
Island, Panama).

1931 Eleutherodactylus lutosus molinoi: Dunn, Occ. Pap. Boston Soc. Nat.
Hist. 5:411.

1952 Eleutherodactylus ridens: Taylor, Univ. Kansas Sci. Bull. 35(5):690,
Fig. 18.

Holotype.—Lost; Nicaragua: Rio San Juan.

This species was originally described from Atlantic versant Nicaragua
near the boundary with Costa Rica. The original account Cope (1867) em-
phasized the areolate venter, reddish dorsal color and dark supratympanic
mark, characteristic of individuals associated with this name. Taylor
(1952:690) subsequently was the first to apply the name ridens to this form,
since Costa Rica examples agreed with Cope’s original account. My ex-
amination of Honduras and Nicaragua specimens confirms their conspeci-
ficity with Costa Rica examples. Although the type is lost, the brief type
description cannot be associated with any other species and I follow Tay-
lor’s use of the name E. ridens for this form.

Barbour’s (1928) Syrrhopus molinoi is based on a 19 mm female that
agrees in detail with Costa Rica specimens in: lacking a well developed heel
tubercle, and having one to several enlarged pointed supraocular tubercles
and the finger disks expanded and rounded. Lynch (1980) concurs with this
allocation based on an independent study of the situation and the holotype
of molinoi. Dunn’s (1931) placement of the latter as a subspecies of lutosus
is clearly incorrect as may be seen from the account of cruentus, above,
and the key presented below. Although Taylor (1952) recognized ridens in
Costa Rica he failed to associate it with the Panamanian frogs called moli-
noi. Myers and Rand (1969) continued to use molinoi for this form on Barro
Colorado Island as a result. Of the three species treated in this paper, ce-
rasinus and ridens occur on Barro Colorado Island; all three occur in the
Panama Canal Zone.

Eleutherodactylus ridens may be separated from other Central American
species in the genus with which it might be confused (except cerasinus and
cruentus) as follows, with features for ridens in parentheses: E. caryophyl-
laceus, E. melanostictus, and E. taeniatus have a well developed heel
tubercle (absent); and E. altae and pardalis have the groin and anterior
thigh marked with large light spots (no spots).

Eleutherodactylus ridens has the following known geographic range: the
Atlantic lowlands of extreme eastern Honduras, Nicaragua, Costa Rica, and
Panama; premontane evergreen forests of Costa Rica and Panama on both
Atlantic and Pacific versants; lowlands of southwestern Costa Rica, and in
the Pacific versant evergreen forests of Panama, western Colombia, and
northwestern Ecuador (15-1200 m).

VOLUME 94, NUMBER 2 419

The following key will distinguish the three species discussed in this re-
port.

1. A distinct well developed pointed to flap-like heel tubercle (cal-
Caine POMOS: ROR SIR a BOA CALS) DR eR, AO EMO EY Oo) he i
— Heel smooth or with a few small warts all of about the same size
and similar to others scattered over upper surface of hind limb; disks
round; surface of upper eyelid with one to several enlarged pointed
supraocular tubercles; anterior and posterior surfaces of thigh, calf
and feet red in life; no nuptial pads; but vocal slits present in adult
males; males to 19 mm, females to 25mm .. Eleutherodactylus ridens
2. A distinct, enlarged, pointed or flap-like supraocular or superciliary
tubercle; subarticular tubercles under fingers and toes not pungent
nor pointed, although often raised; tympanum indistinct (females) or
completely hidden (males) under skin; disks truncate; usually one
to several bright golden-yellow to orange spots in groin and on front
of thigh in life, each outlined by dark pigment; often a series of
oblique black flank bars; nuptial thumb pads but no vocal slits in
adult males; males to 28 mm, females to 42 mm ................
re es ale cee sete Eleutherodactylus cruentus
— No distinct enlarged supraocular nor superciliary tubercle, although
a series of small superciliary tubercles usually present along margin
of eyelid and several small warts on upper eyelid surface; some
subarticular tubercles, at least under toes IIJ-IV, pungent or point-
ed; tympanum thin, distinct; disks truncate; posterior thigh surface
brown with red in groin, on anterior and posterior thigh surfaces and
underside of calf, in life; no discrete dark and light markings in groin;
no nuptial thumb pads but vocal slits present in adult males; males
toms mimi temales tO. 55 Mnl .-..0-.-- .- Eleutherodactylus cerasinus

Acknowledgments

I wish to thank especially the following individuals who have loaned type-
material and been most patient in extending long-term loans of critical ma-
terial: William E. Duellman, University of Kansas (KU); W. Ron Heyer
and the late James A. Peters, U.S. National Museum (USNM); Gunther
Peters, Berlin Museum; Wilmer W. Tanner, Brigham Young University
(BYU); Ernest E. Williams, Harvard College (MCZ). Abbreviations in pa-
rentheses refer to catalogued specimens when mentioned in the text.

Literature Cited

Barbour, T. 1928. New Central American frogs.—Proc. New Engl. Zool. Cl. 10:25-31, pls. 3.
,and E.R. Dunn. 1921. Herpetological novelties —Proc. Biol. Soc. Wash. 34:157-162.

420 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Cope, E. D. 1867. Fourth contribution to the herpetology of tropical America.—Proc. Acad.

Nat. Sci. Phila. 18:123-132.

. 1875. On the Batrachia and Reptilia of Costa Rica.—J. Acad. Nat. Sci. Phila. (letter-

press) ser. 2, (8):93—157.

Dunn, E. R. 1931. New frogs from Panama and Costa Rica.—Occ. Paps. Boston Soc. Nat.

Hist. 5:385-401.

. 1933. Amphibians and reptiles from El Valle de Anton, Panama.—Occ. Paps. Boston

Soc. Nat. Hist. 8:65-79.

Lynch, J. D. 1964. A small collection of anuran amphibians from Panama, with the description

of two new species of Eleutherodactylus (Leptodactylidae).—J. Ohio Herp. Soc.

4(3):65—68. 3 text-figs.

. 1979. The amphibians of the lowland tropical forests.—Univ. Kansas Mus. Nat. Hist.

Monogr. 7:189-215.

. 1980. Systematic status and distribution of some poorly known frogs of the genus

Eleutherodactylus from the Chocoan lowlands of South America.—Herp. 36(2):175-—

189.

Myers, C. W., and A. S. Rand. 1969. Checklist of amphibians and reptiles of Barro Colorado
Island, Panama, with comments of faunal change and sampling.—Smithson. Contr. Zool.
10:1-11.

Peters, W. 1873. Uber eine neue Schildkrotenart, Cinosternun Effeldtii und einige andere
neue oder weniger bekannte Amphibien.—Mon. K. Akad. Wiss. Berlin 1873:603—618.

Savage, J. M. 1965. A new bromeliad frog of the genus Eleutherodactylus from Costa Rica.—
Bull. So. Calif. Acad. Sci. 64(2): 106-110.

Savage, J. M. 1970. On the trail of the golden frog; with Warszewicz and Gabb in Central
America.—Proc. Calif. Acad. Sci. 38(14):273-388.

Taylor, E. H. 1952. A review of the frogs and toads of Costa Rica.—Univ. Kansas Sci. Bull.
35(5):577-942.

Zetek, J., and A. Wetmore. 1951. Report on the Canal Biological Area.—Ann. Rept. Smith-
son. Inst. 1950, App. 10:133-144.

Department of Biological Sciences and Allan Hancock Foundation, Uni-
versity of Southern California, Los Angeles, California 90007.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 421-422

PFLUGELLA, NEW NAME FOR TRICELLARIA
PFLUG, 1965 (MICROPROBLEMATICA), NON
FLEMING, 1828 (BRYOZOA)

Richard W. Huddleston and Drew Haman

Pflug (1965) established the new genus Tricellaria (type-species T. dely-
lensis Pflug) from the Precambrian (Algonkian) of Montana. Pflug (1965: 16)
commented on the possible foraminiferal nature of this form and further
added (1965:23) *‘A problematic fossil is Tricellaria delylensis n. gen. n. sp.
which exhibits several structures similar to Foraminifera.’’ Edwards and
Vevers (1975:347) listed Tricellaria Pflug under Rhizopoda, Protozoa, as did
Tobias (1968:191) who also noted Tricellaria Pflug as *‘a foraminiferan-like
structure.”’

The generic name Tricellaria, however was previously used by Fleming
(1828:502) for a new genus of Recent scrupocellarid bryozoan (type-species
Cellaria ternata Elliss, 1786).

Despite the qustionable foraminifera! affinity of Tricellaria Pflug as well
as its possible removal from the animal kingdom, the taxon is nevertheless
subject to the precepts of the International Code of Zoological Nomencla-
ture (ICZN, Art. 2b). Tricellaria Pflug is a junior homonym of Tricellaria
Fleming and must be replaced (ICZN, Art. 53).

The new name Pflugella is proposed to replace Tricellaria Pflug, 1965,
non Tricellaria Fleming, 1828.

The name Pflugella is constructed in honor of Prof. Dr. Ing. H. D. Pflug,
(Geologisch-Palaontologisches Institut, Liebig University, Giessen) for his
contributions to micropaleontology.

Acknowledgments

We thank Dr. H. D. Pflug for permission to effect correction of the junior
homonym; W. S. Drugg, Chevron Oil Field Research Company for review
of this note, and Chevron Oil Field Research Company for publication per-
mission.

Literature Cited

Edwards, M. A., and H. G. Vevers. 1975. Nomenclator Zoologicus, vol. VII, 1956-1965.—
Zool. Soc. London, 374 pp.

Fleming, J. 1828. History of British Animals. 502+ pp.

International Code of Zoological Nomenclature, 1964.—Intern. Trust Zool. Nomencl. Lon-
don. 176 pp.

422 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Pflug, H. D. 1965. Organische Reste aus der Belt-serie (Algonkium) von Nordamerika.—
Palaeont. Zeitschr. 39:10-25.

Tobias, M. 1968. The Zoological Record, 1965, vol. 102, Sec. 2, Protozoa, pt. II:115—196.

Chevron Oil Field Research Company, P.O. Box 446, La Habra, Califor-
nia 90631.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 423-426

HETEROPTERA RECENTLY COLLECTED IN THE
RAY MOUNTAINS IN ALASKA

Richard C. Froeschner and Libby Halpin

Abstract.—This report on a collection of Heteroptera from the Ray Moun-
tains in Alaska includes a brief description of the areas collected, and con-
tains a list of 18 species (in seven families), including 4 apparent “‘first
records’’ for the State.

The Ray Mountains are located in interior Alaska, approximately 240 km
(150 mi) northwest of Fairbanks and 80 km (50 mi) south of the Arctic Circle.
The range is situated between the Yukon and Koyukuk Rivers. It trends
northwest and is roughly 105 km long (65 mi) and 35 km wide (22 mi). The
topography is diverse with most of the area between 457-1219 m (1500-
4000 ft) elevation though several peaks exceed 1544 m (5000 ft) including
Mount Tozi, the highest at 1682 m (5519 ft).

The range is well characterized as an ‘‘island’’ of subarctic alpine tundra
surrounded by a ‘‘sea’’ of boreal forest. The study area was defined roughly
to include all of the area above treeline (generally near 610 m [2000 ft]) as
well as the forested floodplains and their adjacent forested slopes that finger
up into the range.

The vegetation of the range represents a variety of arctic and alpine, and
boreal forest communities. On floodplains of major streams open evergreen,
deciduous, and mixed forests are present. At elevations above, treeline
forests give way to willow scrub communities. Over the rolling lowlands of
the range, tussock graminoid meadow and dwarf scrub tussock-graminoid
meadow communities dominate. In the higher valleys, dwarf scrub com-
munities are most prevalent, while in the highest parts of the range prostrate
scrubs and barrens predominate (Brock and Burke, 1980; Schubert, 1980).
See Table 1 for descriptions of localities in which collections were made.

During the summer of 1979 L. Halpin collected 39 specimens of terrestrial
Heteroptera (including four unidentifiable nymphs of the family Miridae) in
a study area in the Ray Mountains of Alaska. This collection of 18 species
is of more than casual interest because it includes four species (indicated
by asterisks) for which we have found no previous Alaskan records.

This paper treats the Heteroptera collected during a study of the Ray
Mountains, Alaska, under National Science Foundation Grant No. SPI-
795346.

424 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Description of insect collecting localities.

Average Latitude, Ecosystem Collecting dates
Locality elev. longitude type(s) (all in 1979)
Crash Creek 580 m 65°39'N Lowland tundra Aug. 12-13
1900 ft 150°52’W
Kilo Hot Springs 488 m 65°49'N Boreal forest Aug. 4-7
1600 ft 151°14’W
Lake 1485 452 m 65°52'N Lowland tundra July 18; Aug. 1
1485 ft 151°24’'W
Lower Torment Cr. 580 m 65°49'N Lowland tundra June 24—July 26;
1900 ft 151°26’W Boreal forest July 19-23; Aug. 2-3
Moose Cr. Hdwtrs. 610 m 65°36'N Lowland tundra Aug. 15
2000 ft 150°40’ W Boreal forest
Mt. Tozi Summit 1683 m 65°41'N Alpine barren Aug. 11
5519 ft 150°56’W
Spooky Valley 763 m 65°43'N Upland tundra July 8-11; Aug. 8-9
2500 ft 151°13’W Alpine barren
Tozitna River 580 m 65°38'N Boreal forest Aug. 14
1900 ft 150°46’ W
Twilight Cr. Trib. 824 m 65°40'N Upland/lowland Aug. 11
2700 ft 150°56’W tundra ecotone
Upper Torment Cr. 854 m 65°46’N Upland tundra July 24-30
2800 ft 151°31’W Alpine barren
Acanthosomatidae

*Elasmucha lateralis (Say). Lower Torment Creek, July 29, 1 ¢; Moose
Creek Headwaters, August 15, 1 d, 1 2, 2 Sth instar nymphs. Adults and
nymphs are strongly infuscated, one male so much so that it appears
almost totally black to the unaided eye. Several apparently previously
unreported specimens from Fairbanks, Alaska, are in the National Mu-
seum of Natural History.

Anthocoridae

Anthocoris melanocerus Reuter. Spooky Valley, August 8, 1 3d; Tozitna,
August 14, 1 @.

Lygaeidae

*Geocoris bullatus (Say). Moose Creek Headwaters, August 15, 1 ¢&.
Widely distributed in North America, this species was previously cata-
loged as far northwest as the Yukon in Canada.

Ligyrocoris sylvestris (Linnaeus). Kilo Hot Springs, August 16, 1 9.

VOLUME 94, NUMBER 2 425

Miridae

Calocoris fulvomaculatus (De Geer). Crash Creek, August 12, 1 ¢; Lower
Torment Creek, August 3, 1 6.

Lygocoris contaminatus (Fallen). Lake 1485, July 18, 2 2 2 ; Lower Torment
Creek, June 29, | 6; Kilo Hot Springs, August 13, 1 &.

Lygus columbiensis Knight. Mt. Tozi Summit, August 11, 1 ¢, 1 2 ; Twilight
Creek Tributary, August 11, 1 6.

Lygus varius Knight. Moose Creek Headwaters, August 15, 1 6; Twilight
Creek Tributary, August 11,1 6.

Mecomma gilvipes (Stal). Kilo Hot Springs, August 5, 1 6.

Orthotylus katmai Knight. Twilight Creek Tributary, August 11, 1 @.

Plagiognathus obscurus Uhler. Lower Current Creek, August 3, 1 2.

Polymerus unifasciatus (Fabricius). Tozitna River, August 14, 1 2. This is
the color form with the entire clavus and much of the corium blackened.

Trigonotylus ruficornis (Geoffroy). Twilight Creek Tributary, August 11,
Or

Teratocoris caricis Kirkaldy. Twilight Creek Tributary, August 11,1 ¢&.

Nabidae

*Dolichonabis americolimbatus (Carayon). Lake 1485, August 1, 1 d, 2
22, 1 nymph, latter determined by association with adults. These new
records partially fill the gap between the formerly known North American
range and the disjunct occurrence in eastern Asia as reported by Kerzhner
(1969:521). Such a range suggests a natural migration through Beringia—
whether eastward or westward is not yet clear.

Saldidae

Teloleuca pellucens (Fabricius). Lower Torment Creek, June 29, 1 @.

Scutelleridae

*“Homaemus aeneifrons (Say). Lower Torment Creek, June 29, 1 2; July
29, | nymph. Absence of a male prevents confident subspecies determi-
nation of these specimens, but they quite probably represent Walley’s
(1929:256) western subspecies H. aeneifrons extensus which was origi-
nally described from western North America from Canadian Northwest
Territories south to Colorado.

Literature Cited

Brock, S., and I. Burke. 1980. Vegetation.—IJn N. Farquhar and J. Schubert (eds.), Ray
Mountains, Central Alaska: Environmental Analysis and Resources Statement, Middle-
bury College Press. (In press.)

426 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Kerzhner, I. M. 1969. New and little known Palearctic bugs of the family Nabidae (Heter-
optera).—Entom. Review 47(4):517—525 [English Translation from the Russian, Entom.
Oboz. 47(4):848-863].

Schubert, J. 1980. Ecosystems.—Jn N. Farquhar and J. Schubert (eds.), Ray Mountains,

Central Alaska: Environmental Analysis and Resources Statement, Middlebury College
Press. (In press.)

Walley, G. S. 1929. Notes on Homaemus with a key to the species (Hemip., Scutelleridae).—
Canadian Entom. 61(11):253-256.

(RCF) Department of Entomology, Smithsonian Institution, Washington,
D.C. 20560; (LH) United States Fish and Wildlife Western Alaska Ecolog-
ical Services, 733 W. 4th Ave., Suite 101, Anchorage, Alaska 99501.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 427-436

CHRIOLEPIS VESPA, A NEW SPECIES OF GOBIID
FISH FROM THE NORTHEASTERN
GULF OF MEXICO

Philip A. Hastings and Stephen A. Bortone

Abstract.—Chriolepis vespa is described from the northeastern Gulf of
Mexico. It is distinguished from the similar Chriolepis benthonis Ginsburg
by its greater number of second dorsal-fin elements (10 vs. 9), greater degree
of scale spination, longer pectoral fin, longer fourth and shorter fifth pelvic
rays, tongue shape, and color pattern. The degree of branching of pelvic-fin
rays one through four increases with growth in C. vespa. Varicus Robins
and Bohlke has been distinguished from Chriolepis Gilbert on the basis of
the degree of branching of the pelvic-fin rays (unbranched in Varicus) and
other characters which prove to be inconsistent since the discovery of C.
vespa. The systematics of this group of seven-spined gobies which lack
sensory pores is in need of revision.

The gobiid fish genus Chriolepis Gilbert 1892, is presently poorly known
in the western Atlantic. Only two species, Chriolepis fisheri Herre 1942,
and Chriolepis benthonis Ginsburg 1953, have been described, each from
a single specimen. Recent collecting on the offshore continental shelf in the
northeastern Gulf of Mexico produced a series of specimens of a new goby
referable to Chriolepis. Examination of other western Atlantic specimens
of this genus indicates the presence of several other undescribed species,
whose descriptions await the accumulation of additional specimens.

Methods.—Al\l measurements were made with dial calipers to the nearest
0.1 mm. Meristic and morphometric data were recorded following the pro-
cedures of Bohlke and Robins (1968) with the following emendations and
additions: fleshy interorbital width is the least distance between the fleshy
portions of the supraorbital area; head depth at occiput is the vertical dis-
tance from the midline of the occiput to the ventral contour of the head;
head depth at preoperculum is the vertical distance from the midline above
the posterior edge of the preoperculum to the ventral contour of the head;
head width is the maximum horizontal distance between the opercula; in-
termandibular width is the distance between the posterior mandible tips;
body depth at anal-fin origin is the vertical distance from the anal-fin origin
to the dorsal body contour; pelvic-fin spine length is the distance from the
Spine insertion to its tip; pelvic-fin ray length is the distance from the ray
insertion to the tip of the longest branch.

428 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Photograph of holotype of Chriolepis vespa, USNM 221523, 26.6 mm SL, female.

Institutional abbreviations are as follows: GCRL, Gulf Coast Research
Laboratory, Ocean Springs, Mississippi; SU, Stanford University (collec-
tions now deposited at the California Academy of Sciences, San Francisco,
California); USNM, United States National Museum of Natural History,
Washington, D.C.; UF, Florida State Museum, University of Florida,
Gainesville; UMML, University of Miami, Rosenstiel School of Marine and
Atmospheric Sciences, Miami, Florida.

Chriolepis vespa, new species
Wasp goby
Pigs. 2.3

Holotype.—USNM 221523, 26.6 mm SL female, northeastern Gulf of
Mexico, 28°24’N, 85°15’'W, R/V BELLOWS (cruise Bureau of Land Man-
agement, BLM 15, station III-C), collected with a 6.1 m semi-balloon trawl
on 26 July 1975, depth 183 m, bottom composed of gray mud.

Paratypes.—USNM 221524, 7 (31.4-17.9 mm SL), collected with holo-
type; UF 28037, 1 (26.4 mm SL), 27°50'N, 84°42’W (R/V BELLOWS cruise
BLM 15, station II-C), 27 July 1975, 183 m; UF 28038, 1 (34.1 mm SL),

Fig. 2. Drawing of holotype of Chriolepis vespa.

VOLUME 94, NUMBER 2 429

2mm

rt

secs menRelinela))) 222 naam

ee ee

SSS
vanes) NLL ao oo noe we—
\((ansaonDD Danae
te oor uesSesce s eoeeeee ee
LTT VTj

SOC ee
“SSS0a55

ITT —,

=

Fig. 3. Left pelvic fins of Chriolepis vespa. A, USNM 221524, 19.5 mm SL; B, UF 28038,
34.1 mm SL.

28°24'N, 85°15’ W (R/V BELLOWS cruise BLM 13, station III-C), 29 August
1977, 183 m; GCRL 16972 (previously USAIC 05983), 1 (24.2 mm SL), north-
eastern Gulf of Mexico, 29°45.6’N, 87°46.2'’W, 7 Feb. 1978, 35 m.

Diagnosis.—A plump-headed species of Chriolepis with 10 (I, 9) second
dorsal-fin elements and 15-17 pectoral rays. Degree of branching of first
through fourth pelvic-fin rays increasing with growth. Fifth pelvic-fin ray
unbranched and short. Two to six rows of cycloid scales followed by six to
nine rows of ctenoid scales on posterior third of body. Body pale yellowish
with four olive bars with intervening olive saddles. Head with a yellow
subocular bar in life.

Description.—Meristics and morphometrics of the type specimens are
given below and in Table 1. Dorsal VII-10; anal 8 (7-9); pectoral 16 (15-
17); vertebrae 27 (11 trunk + 16 caudal).

First through fifth spines of first dorsal fin closely spaced, sixth and sev-
enth more widely spaced, none filamentous. Dorsal fins separate though
closely spaced. First element of second dorsal fin a slender spine followed
by nine branched rays (last two rays counted as one). First anal-fin element
a slender spine followed by 6-8 (typically 7 or 8) branched rays (last two
counted as one). Caudal fin truncate in largest specimens, rounded in others.
Caudal fin composed of 16-17 total segmented rays (8-9 upper + 7-8 lower
segmented rays); not all segmented rays are branched. Upper and lower

430 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Meristics and morphometrics (expressed as thousandths of the standard length)
of Chriolepis vespa.

Paratypes
Holotype Range Mean
Meristics
First dorsal-fin elements VII VII VII
Second dorsal-fin elements 10 10 10
Anal-fin elements 8 7-9 8.2
Pectoral-fin rays 15 + 16 15-17 16.0
Morphometrics
Standard length (mm) 26.6 17.9-34.1 MSL
Head length 305 293-354 327
Snout length 56 52-85 65
Maximum eye diameter 105 98-125 108
Interorbital width 9 8-15 11
Postorbital distance 162 147-192 169
Head depth at occiput N32 120-185 147
Head depth at preoperculum 165 155-226 167
Head width 214 196-273 232
Upper jaw length 109 111-123 116
Intermandibular width 132 112-170 149
Body depth at first dorsal-fin origin 180 130-187 163
Body depth at anal-fin origin 158 140-169 154
Caudal-peduncle depth 113 107-118 112
Caudal-fin length 263 265-308 280
Pectoral-fin length 323 284-364 322
Pelvic fin
Spine length 124 110-128 115
Fourth ray length 293 271-365 312
Fifth ray length 90 41-87 Ui
Pelvic-fin base to anal-fin origin distance 327 282-371 327

caudal fin composed of 7-8 and 6-8 branched rays, respectively (13-16 total
branched rays). Pectoral fin angulate (central rays longest), extending pos-
teriorly to anal-fin origin when adpressed. Pelvic fin I, 5. Pelvic spine short
(11-13% of SL). First through fourth pelvic rays increasing in length, the
fourth being longest and extending past anus when adpressed (but falling
short of the anal-fin origin). Fifth ray short, inconspicuous, adhering closely
to the fourth and never branched (Fig. 3). First through fourth rays flattened
slightly distally and branched in all specimens examined (Fig. 3); the extent
of branching increases with fish length. In smaller individuals (Fig. 3A, 19.5
mm SL), the first through fourth rays with only one shallow branch. The
largest specimen (Fig. 3B, 34.1 mm SL) with highly branched rays. In larger
specimens, first ray with a secondary bifurcation near the tip of the medial
branch of the primary bifurcation. Second, third, and fourth rays with both

VOLUME 94, NUMBER 2 431

secondary and tertiary bifurcations. Typically secondary and tertiary bifur-
cations occur on the medial branch of the preceding bifurcation. Specimens
of intermediate size show intermediate patterns of pelvic fin branching.

Pelvic-fin origin under pectoral-fin origin. Pelvic fins separate, with no
interspinous membrane and no membrane connecting the fins medially;
however, an inconspicuous low fleshy ridge between the inner bases of the
pelvic fins is evident in some specimens.

Head plump, approximately one-third of SL. Eye large (maximum di-
ameter about one-third of head length), slightly elevated and superior. In-
terorbit narrow. Frontals reduced to vestiges between orbits. Mouth ter-
minal, inclined upward approximately 30° from the horizontal. Upper jaw
extends posteriorly to below anterior edge of pupil. No head pores. Sensory
papillae in rows between nostrils, on preorbit, suborbit, cheek, operculum,
lower jaw symphysis, posterior edge of branchiostegal membrane, above
pectoral-fin base, and along lateral midline of body anterior to scale rows
(Fig. 2). Anterior nostril with a short tube, about one-half pupil diameter in
length. Posterior nostril with a minute tube, about one-fourth length of an-
terior nostril tube. Tongue shallowly indented. Buccal valves well devel-
oped.

Posterior portion of body scaled. Scales somewhat caducous. Total scale
rows 9-14: 2-6 rows of cycloid scales anteriorly; 6—9 rows of ctenoid scales
posteriorly. Origin of cycloid scale rows variable, from under the fifth to the
ninth element of the second dorsal fin (sixth in holotype). Origin of ctenoid
scales also variable, from under seventh to below last element of second
dorsal fin (tenth in holotype). Four enlarged basicaudal ctenoid scales pres-
ent on upper and lower caudal fin, one upper and one lower on each side.
Basicaudal scales missing from left side of holotype (Fig. 2). Ctenii on each
basicaudal scale large, conspicuous, and variable in number (4-12). A few
additional ctenoid scales on caudal fin between basicaudal scales.

Teeth in lower jaw in four rows anteriorly. Outermost row composed of
widely-spaced, enlarged recurved canines. Innermost row composed of
more closely-spaced, slightly smaller recurved canines. Inner and outer
rows separated anteriorly by two rows of irregular low-pointed teeth. Lat-
erally the outermost and central rows are lacking. Only the innermost row
of canines present at angle of jaw. Teeth in upper jaw in about five rows
anteriorly, outermost row composed of widely-spaced, enlarged recurved
canines; four rows of irregular low-pointed teeth medial to outer row with
a few slightly enlarged teeth irregularly spaced within medial rows. Inner-
most row close-set, with teeth not enlarged as in lower jaw. Outermost row
extends to angle of jaw and tooth size decreases posteriorly. Inner rows
decrease in number and size posteriorly until only two irregular rows of low
teeth remain at angle of jaw. Modified or enlarged lateral canines lacking in
either jaw. Teeth lacking on vomer and palatines.

Color notes taken from a fresh paratype (USNM 221524, 31.4 mm SL)

432 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

supplemented by notes taken from a kodachrome of the same specimen are
as follows: body pale yellowish, with pinkish tint anteriorly near pectoral
base. Body translucent posteriorly (vertebral column visible through caudal
peduncle). Four prominent olive, vertical bars on body; first bar under mid-
spinous dorsal fin, second just posterior to second dorsal-fin origin, third
connecting end of second dorsal fin with end of anal fin, fourth at caudal
base. All bars extend to ventral outline of body and increase in intensity
posteriorly. Two olive saddles on nape, first above preoperculum, second
just anterior to pectoral-fin base. Four additional olive saddles on body, first
at spinous dorsal-fin origin and one between each of the vertical bars; all
saddles paler than vertical bars, and not extending below midline of body.
Opercular region, lateral pectoral base, and chest cream-colored with a pink
cast. Remaining body white ventrally. Head white from snout to preoper-
culum. Nape with an olive cast. Faint pink-orange pigment present on oper-
culum. Bright yellow subocular bar extending vertically from orbit to angle
of jaw. Lower jaw white. Branchiostegal membranes yellow. Iris golden.
Spinous dorsal fin with a wide, yellow-olive band medially and a white band
distally. Distal tips of spines yellow-olive. Second dorsal fin with three rows
of yellow-olive spots separated by three white bands (yellow-olive proxi-
mally followed distally by alternating white and yellow-olive bands). Second
dorsal-fin margin black. Anal fin yellow-olive medially, black distally. Distal
black band occupies one-fifth of fin anteriorly and grades to occupy three-
fifths of the fin posteriorly. Two vertical yellow bars on caudal fin followed
distally by a white bar. Distal margin of fin with a black band. Lower caudal-
fin rays pale yellow with sparsely scattered chromatophores. Pectoral fin
yellow-orange. Pelvic fin yellow-orange proximally, gray distally.

Color of preserved (45% isopropynol) specimens (Figs. 1, 2) as follows:
body pallid. Vertical bars and saddles clearly visible on body as clusters of
chromatophores. Few chromatophores scattered elsewhere over dorsal and
lateral aspects of body. Abdomen, thorax, ventral portion of lateral pectoral-
fin base, branchiostegal membranes, and isthmus unpigmented. Nape with
contracted chromatophores. Sparse chromatophores present on cheek, sub-
orbit, preorbit, interorbit, and along upper and lower jaws. Iris black. Spi-
nous dorsal fin with scattered chromatophores, most dense at tips of spines.
Second dorsal fin with scattered chromatophores, most dense on distal mar-
gin. Anal fin unpigmented proximally, but with densely-spaced chromato-
phores on distal margin. Caudal fin with few scattered chromatophores cen-
trally but with closely-spaced chromatophores peripherally on posterior and
ventral margins. Upper portion of outer pectoral-fin base with chromato-
phores forming an irregular blotch. Few chromatophores on pectoral fin.
Pelvic fin with few scattered chromatophores.

Comparisons.—Chriolepis vespa morphologically resembles C. benthonis
Ginsburg 1953, described from a single specimen from off the Yucatan Pen-

VOLUME 94, NUMBER 2 433

insula in 154 m of water. The two species are similar in overall head and
body shape, number of anal-fin rays (8 in C. benthonis), and pectoral-fin
rays (16 in C. benthonis). They differ in number of second dorsal-fin rays
(9 in C. benthonis) and in squamation. Although several scales are missing
from the holotype of C. benthonis, salient differences are apparent in those
that remain. In C. benthonis there are seven enlarged scales located mid-
laterally on the caudal peduncle. Only the last three or four of these possess
a few weakly developed ctenii. Small cycloid scales are present on the
caudal peduncle above and below the central seven ctenoid scales. Cycloid
scales also extend forward in a wedge to under the sixth element of the
second dorsal fin. A few cycloid scales are present on the caudal-fin base.
No enlarged basicaudal ctenoid scales are present, although these could
have been lost. Ginsburg (1953) stated that modified scales were present on
the caudal base but most were missing. In C. vespa, six to nine rows of
well-developed ctenoid scales are present on the body posteriorly. Further,
most scales above and below the mid-lateral series possess well-developed
ctenii (as many as twelve on one scale), while the only cycloid scales present
are those anterior to the mid-lateral ctenoid rows. The ctenii on most scales
of C. vespa are larger than any of those that remain on the holotype of C.
benthonis. In C. vespa a few ctenoid scales are present on the caudal fin
between the modified basicaudal scales. Chriolepis vespa specimens and
the holotype of C. benthonis also differ in some body proportions: C. ben-
thonis has a shorter pectoral fin (248 thousandths of the SL), a shorter fourth
pelvic-fin ray (247), and a longer fifth pelvic ray (105). Further, the two
species differ in tongue shape (rounded in C. benthonis, indented in C.
vespa).

R. S. Birdsong kindly allowed examination of a specimen of Chriolepis
from Arrowsmith Bank off the Yucatan Peninsula (UMML uncat., R/V
PILLSBURY station 584) which we believe represents the second known
specimen of C. benthonis. It agrees with the holotype in dorsal-fin elements
(VII-9) and anal-fin elements (8) and was taken relatively near (310 km SE)
the type locality though in deeper water (approximately 350 vs. 154 m).
Unfortunately, the scales are missing from the PILLSBURY specimen. A
kodachrome taken of this specimen when initially captured reveals a pale
yellow body and a vivid yellow wash on the head. No vertical bars are
present on the body, although the head and body do have scattered chro-
matophores. Ginsburg (1953:22) described the color of the holotype of C.
benthonis as *‘. .. a rather uniform light yellowish, probably faded .. .”’
and thus similar to the PILLSBURY specimen. If this uniform yellow color
reflects the true coloration of C. benthonis and is not the result of fading,
then C. vespa and C. benthonis differ markedly in color pattern, as vertical
bars are prominent in C. vespa. Recent examination of the holotype of C.
benthonis revealed no evidence of vertical bars.

434 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

According to the original description of the holotype (Herre, 1942), C.
fisheri differs from C. vespa in several characters including head shape
(more flattened in C. fisheri) and degree of squamation (body naked in C.
fisheri with only one upper and one lower basicaudal scale present). Me-
ristic differences between the two species include the number of anal-fin
elements (10 in C. fisheri) and the number of second dorsal-fin elements (11
in C. fisheri), though a problematic specimen of C. vespa has 11 second
dorsal-fin elements (see below). Further, the two species differ in coloration;
Six narrow crossbars, some of which extend upon the dorsal and anal fins,
are present on the body of C. fisheri (Herre, 1942) while four are confined
to the body of C. vespa.

Chriolepis vespa differs from the somewhat similar Varicus bucca Robins
and Bohlke 1961, principally in the degree of branching of the pelvic-fin rays
(unbranched in V. bucca), and in having fewer scale rows (27 in V. bucca).
Chriolepis vespa differs from Varicus marilynae Gilmore 1979, which has
slightly bifurcated pelvic-fin rays, in the number of second dorsal-fin ele-
ments (9 in V. marilynae), in having fewer scale rows (18-19 in V. mari-
lynae), and in coloration (lower portion of head orange in V. marilynae).

Distribution and habitat.—The type specimens are from the northeastern
Gulf of Mexico from off Mobile Bay, Alabama, to off Tampa Bay, Florida.
One specimen was taken in 35 m while all others were from 183 m. Substrate
at all localities was grey mud. An additional specimen which may be refer-
able to this species (see below) was taken off Guyana (7°10’N, 53°36’ W).

Etymology.—The name vespa is from the Latin “‘vespa,’’ meaning wasp.
This name is chosen to bring attention to the wasp-like olive banding of this
species.

Discussion.—An additional specimen from off Guyana (UMML uncata-
logued, R/V PILLSBURY station 658) which may represent C. vespa has ap-
parently been lost (R. S. Birdsong and C. R. Robins, pers. comm.). A descrip-
tion of this specimen based on a black/white photograph and radiograph
provided by Birdsong follows. Vertebrae 27; dorsal VII-11; anal 9. Ctenoid
scales present on lateral portion of caudal peduncle. Enlarged ctenoid basi-
caudal scales (one upper and one lower) present on caudal-fin base. Chromato-
phores scattered over head and body, concentrated in four vertical bars on
body. Light subocular bar present. Anal-fin margin and lower and distal
caudal-fin margins black.

C. R. Robins, who examined this specimen and a kodachrome of a fresh
paratype of C. vespa, indicated (pers. comm.) that the life colors of the
PILLSBURY specimen closely resembled those of C. vespa. This record
is tentatively referred to C. vespa because of its similar meristics (differs
in having 11 instead of 10 second dorsal-fin elements) and color pattern.
Considering the distance (approximately 3200 km) between collecting sites,
this specimen needs further examination before positive identification can

VOLUME 94, NUMBER 2 435

be made. If it is found and proves to be C. vespa, it would increase the
species’ known distribution considerably.

The genera Chriolepis Gilbert and Varicus Robins and Bohlke have been
distinguished (Bohlke and Robins, 1968) by the following characters: state
of pelvic-fin ray branching (branched in Chriolepis, unbranched in Varicus),
presence or absence of a low fleshy membrane connecting the inner pelvic-
fin rays (absent in Chriolepis, present in Varicus), and tongue shape (round-
ed in Chriolepis, bilobed in Varicus). Generic allocation of C. vespa is
based on the extent of branching of the pelvic-fin rays. However, as the
degree of pelvic-fin ray branching increases ontogenetically in C. vespa,
this character may be of little value in distinguishing these genera. Further,
the other two characters used to separate these genera are invalidated by
the discovery of C. vespa. A low fleshy ridge is present between the inner
pelvic rays of C. vespa (this ridge is also present on several eastern Pacific
members of the genus Chriolepis, L. T. Findley, pers. comm.; Findley,
1975) and the tongue is not rounded but has a distinct indentation anteriorly.

The generic alignment of Varicus marilynae Gilmore 1979, which has
slightly bifurcated pelvic rays, is open to question considering the variation
of this character in C. vespa and the inconsistency of the other available
generic distinctions. The western Atlantic members of this entire group of
seven-spined gobies with divided pelvic fins and no head pores is in need
of revision, as several apparently undescribed forms with both branched
and unbranched pelvic-fin rays exist. Definitive generic allocation of these
and previously described forms awaits further study.

Comparative material examined.—Chriolepis sp., tentatively identified as
Chriolepis vespa: UMML uncatalogued, R/V PILLSBURY station 658,
07°10’N, 53°36’W, off Guyana, 4 July 1968, depth 126-135 m (specimen not
examined, apparently lost, black/white photograph and radiograph exam-
ined). Chriolepis benthonis: USNM 47641, holotype, off the Yucatan Pen-
insula, Mexico; UMML uncatalogued, R/V PILLSBURY station 584, 21°02'N,
86°24’W, Arrowsmith Bank off the Yucatan Peninsula, Mexico, 23 May
1967, depth 347-353 m. Chriolepis fisheri: SU 37262, holotype, Barbados
(radiograph only). Varicus bucca: UMML 7114, paratype, Caribbean Sea
(cleared-and-stained). Varicus marilynae: USNM 218406, holotype, Atlan-
tic Ocean, Florida east coast; UF 24757, paratype, Gulf of Mexico, Florida
west coast.

Acknowledgments

We thank C. R. Robins and R. S. Birdsong for allowing us to describe
this species, which they also recognized as undescribed. Birdsong kindly
provided several specimens and radiographs, a photograph and radiograph
of the apparently lost R/V PILLSBURY specimen, and helpful comments

436 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

on the manuscript. L. T. Findley provided insight into Chriolepis system-
atics and a meticulous review of the manuscript. E. A. Lachner (USNM),
C. R. Robins (UMML), and R. L. Shipp (USAIC) allowed examination of
specimens in their care. T. Smoyer provided photographic assistance and
C. Baker drew Fig. 3. Harbor Branch Foundation, Inc. contribution number
209.

Literature Cited

Bohlke, J. E., and C. R. Robins. 1968. Western Atlantic seven-spined gobies, with descrip-
tions of ten new species and a new genus, and comments on Pacific relatives.—Proc.
Acad. Nat. Sci. Philadelphia 120(3):45-174.

Findley, L. T. 1975. A new species of goby from Malpelo Island (Teleostei: Gobiidae: Chri-
olepis).—Smithsonian Contr. Zool. 176:94—98.

Gilbert, C. H. 1892. Scientific results of explorations by the U.S. Fish Commission steamer
ALBATROSS, XXII: Descriptions of thirty-four new species of fishes collected in 1888
and 1889, principally among the Santa Barbara Islands and in the Gulf of California.—
Proc. U.S. Natl. Mus. 14(880):539-566.

Gilmore, R. G. 1979. Varicus marilynae, a new gobiid fish from Florida.—Copeia 1979(1): 126—
128.

Ginsburg, I. 1953. Ten new American gobioid fishes in the United States National Museum,
including additions to a revision of Gobionellus.—J. Washington Acad. Sci. 43(1):18-
26.

Herre, A. W. C. T. 1942. Notes on a collection of fishes from Antigua and Barbados, British
West Indies.—Stanford Univ. Publ., Biol. Sci. 7(2):285-—305.

Robins, C. R., and J. E. Bohlke. 1961. A new gobioid fish from the Antilles and comments
on Ctenogobius fasciatus and C. curtisi.—Copeia 1961(1):46—50.

(PAH) Harbor Branch Foundation, Inc., Ft. Pierce, Florida. Present ad-
dress: Dept. of Ecology and Evolutionary Biology, University of Arizona,
Tucson, Arizona 85721; (SAB) Dept. of Biology, University of West Flor-
ida, Pensacola, Florida 32504.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 437-444

A NEW GENUS AND A NEW SPECIES OF
HELMINTHOGLYPTID LAND SNAILS FROM
THE MOJAVE DESERT OF CALIFORNIA

Walter B. Miller

Abstract.—A new genus, Eremariontoides, is erected for Eremarionta
argus (Edson, 1912) because its anatomy differs significantly from that of
Eremarionta Pilsbry, 1913. A new species of Eremarionta, E. greggi, is
described for populations that had been considered conspecific with E. ar-
gus but whose anatomy reveals them to be typical Eremarionta.

In 1912, H. M. Edson described Sonorella argus from dead shells col-
lected by A. M. Strong at the Iron Cap copper mine in Revenue Canyon,
Argus Mts., California. Pilsbry (1939) assigned this species to the genus
Micrarionta Ancey, 1880, subgenus Eremarionta Pilsbry, 1913, on the basis
of shell characters only. Eremarionta was raised to generic rank by Be-
quaert and Miller (1973) because its anatomy and shell characters were
sufficiently different from Micrarionta to render questionable the implied
close phylogeny conferred by subgeneric status.

W. O. Gregg and I have dissected topotypes of E. argus and found that
the anatomy of the reproductive system is considerably different from that
of typical Eremarionta. There is no dart sac, no mucus glands, and the
epiphallic caecum is reduced to a minute apical appendage on the epiphallus;
furthermore, the penis and epiphallus are greatly shortened from the usual
Eremarionta structures.

Over a period of years, Gregg and I have obtained specimens of E. argus
from other localities in the Argus Mts., especially Homewood Canyon, at
the southern end of the range, and we have found that all specimens dis-
sected exhibit the same anatomy as the topotypes. More recently, speci-
mens from the Slate Range, a spur of the Argus Range, have also revealed
a similar anatomy.

By contrast, specimens from Johnson Canyon, in the Panamint Mts.,
which had been considered conspecific with E. argus on the basis of shell
characters, have revealed a typical Eremarionta anatomy, with dart sac,
mucus glands, long epiphallic caecum, and appropriate penial characteris-
tics. Additional collecting in the eastern end of the Avawatz Mts., east of
Fort Irwin, has yielded specimens similar to those found in the Panamint
Mts., both in shell morphology and in reproductive anatomy.

It is apparent that the Argus and Slate populations have undergone major

438 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. A, B, C, Shell of Eremariontoides argus; D, E, F, Shell of Eremarionta gregg
holotype. Scale line, 1 cm.

VOLUME 94, NUMBER 2 439

genetic changes from the populations in the Panamint and Avawatz ranges,
probably a form of saltational speciation by chromosomal rearrangement,
along the lines of the probable evolution of Mohavelix micrometalleus (Ber-
ry, 1930) from a population of Helminthoglypta micrometalleoides Miller,
1970. Unfortunately, to date, Noorullah Babrakzai has been unsuccessful
in attempts to stain for C-bands and G-bands on the relatively very small
and numerous chromosomes of any of the Argus, Slate, Panamint, or Ava-
watz populations. Initial investigations on chromosome spreads of the Argus
population, indicate a chromosome number of 2n = 60 (Babrakzai, personal
communication). Babrakzai and I suspect that a comparison of chromosome
bands would reveal the type of chromosomal rearrangement that has prob-
ably occurred.

Meanwhile, in order to reflect correctly the systematic positions of the
different populations, it has become necessary to erect a new genus for E.
argus and to describe a new species for the Panamint and Avawatz popu-
lations which, heretofore, have been considered conspecific with E. argus.

The following abbreviations for repositories of material are employed:
ANSP = Academy of Natural Sciences of Philadelphia; CAS = California
Academy of Sciences; CCC = Carl C. Christensen collection; FMNH =
Field Museum of Natural History; USNM = U.S. National Museum of
Natural History; UTEP = University of Texas at El Paso; WBM = Walter
B. Miller collection.

Eremariontoides, new genus
Figs. 1A, B, C; 2A, B

Description.—Eremariontoides is a helminthoglyptid genus with shells
similar to Eremarionta but with a reproductive anatomy characterized by
the complete absence of dart apparatus and mucus glands. Additionally, the
spermathecal duct and the spermathecal diverticulum are similar to those
of Eremarionta, but the epiphallic caecum is reduced to a minute tip at the
end of the epiphallus. The epiphallus is short and contains 3—4 longitudinal,
anastomosing, thick pilasters which can protrude only slightly to form a
minute, verge-like penis-papilla, into the short, saccular penial chamber.

Type-species.—Eremariontoides argus (Edson, 1912).

Distribution.—Eremariontoides is a monotypic genus which has been con-
firmed only in the Argus and the Slate Ranges of California.

Differential diagnosis.—Eremariontoides joins the list of the west North
American helminthoglyptid genera that have lost the dart apparatus and
mucus glands, namely Sonorelix Berry, 1943, Sonorella Pilsbry, 1900, Mo-
havelix Berry, 1943, Greggelix Miller, 1972, and Tryonigens Pilsbry, 1927.

It is separated from Sonorella and Mohavelix by its substantial sper-
mathecal diverticulum, its foreshortened and thick epiphallus, and its short

440 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

lO mm

Fig. 2. A, Reproductive system of Eremariontoides argus; B, Reproductive system of
Eremarionta greggi. Both figures to same scale; ovotestes omitted. ag, albumen gland; ds, dart
sac; ec, epiphallic caecum; ep, epiphallus; go, genital orifice; hd, hermaphroditic duct; mb,
mucus bulb; mg, mucus gland membrane; pe, penis; pr, penial retractor muscle; pt, prostate;
sd, spermathecal duct; sp, spermatheca; sv, spermathecal diverticulum; ut, uterus; va, vagina;
vd, vas deferens. A, WOG 7765-c stained whole mount, Revenue Canyon, Argus Mts. B,
WBM 5897 stained whole mount, Johnson Canyon, Panamint Mts.

VOLUME 94, NUMBER 2 44]

0.5mm

Meese
ander

24g PPCM Adem ee
ATW Alt

Si ag.
Ae
tae Sad

0.5mm

Fig. 3. A and B. Internal structures of male reproductive system of Eremariontoides argus.
C and D. Internal structures of male reproductive system of Eremarionta greggi. All figures
drawn from stained whole mounts as seen by camera lucida through compound microscope.
A, WBM 5896; B, WOG 7765-c; both from Revenue Canyon, Argus Mts.; C, WBM 5897: D,
WOG 7971-c; both from Johnson Canyon, Panamint Mts.; ec, epiphallic caecum; ep, epiphal-
lus; lec, lumen of epiphallic caecum; lep, lumen of epiphallus; pc, penial chamber; pe, penis;
pi, pilaster; pp, penis papilla; pr, penial retractor muscle; vd, vas deferens; ve, verge.

442 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

penis with occasionally a very slightly everted penis-papilla formed by the
protruding distal ends of epiphallic pilasters. It is separated from Sonorelix
and Greggelix by its very short epiphallic caecum and the absence of a true,
non-invaginable verge. 7ryonigens stands apart from all other genera by its
extremely short spermathecal duct; otherwise, Tryonigens most closely re-
sembles Sonorella, and differs from Eremariontoides by its sizeable, non-
invaginable verge situated in the penial chamber, and by its lack of sper-
mathecal diverticulum.

Eremarionta greggi, new species
Fics, [DME OES 2B. 3)

Description of shell of holotype.—Shell small for the genus, with rounded
whorls and moderately open umbilicus; color pale-brown, with a typical
narrow, dark-brown band above the periphery, bordered on each side by a
paler, whitish band. The body whorl descends sharply to the aperture bor-
dered by a slightly expanded peristome. Embryonic shell of 1'2 whorls
initially glassy smooth, then with light radial wrinkles gradually superim-
posed with spirally arranged, elongate papillae. Beyond the embryonic shell,
the papillae are present for about one more half whorl and then disappear
completely; light radial wrinkles persist on all whorls to the aperture. The
aperture 1s rounded-oval with margins converging to the thin parietal callus;
the columellar margin of the aperture expanded and slightly reflexed over
the umbilicus.

Measurements of holotype.—Shell diameter 12.4 mm, shell height 6.5
mm, unbilicus width 2.1 mm, number of whorls 4.

Reproductive system anatomy of holotype.—The reproductive system is
typical for the genus, with a dart-containing dart sac inserted on the vagina,
2 mucus glands whose glandular tissues descend and spread around the
vagina and whose ducts lead separately from each of the 2 bulbs into the
vagina immediately above the opening of the dart sac. The spermathecal
duct is typical for the genus and bears a moderately long spermathecal
diverticulum. The penial retractor muscle is attached to the epiphallus which
is equipped with a moderately long epiphallic caecum; at its distal end, the
epiphallus consists of a short inner tube within an outer tube. The typical
penial sac contains a verge at its proximal end which is formed by a partial
eversion of the inner tube of the epiphallus.

Remarks.—The shell of Eremarionta greggi is in all respects similar to
that of Eremariontoides argus, with which it has been considered conspe-
cific. As stated by Pilsbry (1939): ‘‘the depressed shape and very wide last
whorl are its most prominent features.’’ Adult shell measurements in the
type lot vary from a minimum diameter of 10.7 mm to a maximum of 13.5
mm and from a minimum height of 5.7 mm to a maximum of 6.9 mm.

VOLUME 94, NUMBER 2 443

The reproductive anatomy is typical for the genus and is not diagnostically
distinctive. The verge varies in length in different specimens depending on
the degree of evagination of the distal inner epiphallic tube into the penial
sac.

I have examined the reproductive system of most of the described species
of Eremarionta and, in all specimens, I have noted a distinct verge formed
by partial eversion of the inner epiphallic tube. Although the anatomy of
this type of eversible verge indicated that a complete retraction should be
possible, I have never observed such complete retraction. It is possible,
however, that the ever-present partial eversion may be an artifact of prep-
aration caused by the method of obtaining the body of the animal by drown-
ing and heating to 60°C in order to avoid breaking the shell. Live, adult
specimens of most species of Eremarionta are usually rare and difficult to
obtain, so that every effort is made to preserve intact both the shell as well
as the body of the animal.

Distribution.—Eremarionta greggi is known only from the Panamint
Mountains and the Avawatz Mountains of California. It may be present in
the Owl Head Mountains and possibly in other nearby ranges where E.
argus had been said to occur; it will be necessary, however, to examine the
anatomy of all such reported specimens before a definite diagnosis can be
made.

Disposition of types.—Holotype, USNM 784583; Paratypes, USNM
784584; ANSP 353858; CAS 60745-60747; CCC 3949; FMNH 198998; UTEP
7958; WBM 5897.

Localities of Collections

Topotypes of Eremariontoides argus were collected from the following
locality: Large, north-facing rockslide located 1.3 miles up Revenue Canyon
from the limestone crusher plant, Argus Mts., Inyo Co., CA.

Additional collections of E. argus were made at the following localities:
1. Homewood Canyon, Argus Mts., in north-facing rockslides. 2. East side
of Slate Mts. in rocks at base of ravines at south end of Panamint Dry Lake.

Type-locality for Eremarionta greggi: Johnson Canyon, Panamint Mts.,
in large north-facing rockslide at a point 6.5 road miles up Johnson Canyon
road from the main west side road.

Populations of E. greggi were also confirmed in the Avawatz Mts., San
Bernardino Co., CA, in north-facing rockslides along the Silver Lake-Fort
Irwin road at the boundary of Fort Irwin.

Acknowledgments

The late Wendell O. Gregg introduced me to desert collecting in 1956. He
was my constant companion and teacher for many years, and he has be-

444 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

queathed to me a large number of undescribed populations of California
land snails. His initial investigations in Eremarionta and Eremariontoides
were responsible for the eventual preparation of this paper. I am also in-
debted for the assistance and camaraderie of Drs. Noorullah Babrakzai,
Carl C. Christensen, and H. Lee Fairbanks in the course of several trips to
field localities cited above. Their enthusiasm and encouragement contrib-
uted significantly to the successful elucidation of this perplexing problem of
separating and identifying the many widespread populations of both species.

Literature Cited

Bequaert, J. C., and W. B. Miller. 1973. The mollusks of the arid Southwest.—University of
Arizona Press: xvi + 271 pp.

Edson, H. 1912. Two new land shells from California.—Nautilus 26(4):37.

Miller, W. B. 1970. A new species of Helminthoglypta from the Mojave Desert.—Veliger
12(3):275—278.

Pilsbry, H. A. 1939. Land Mollusca of North America (north of Mexico).—Acad. Nat. Sci.
Monogr. 3 (Vol. I, Pt. 1): xvii + 573 + 1-ix pp.

Department of General Biology, University of Arizona, Tucson, Arizona
SSpZile

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 445-449

A NEW SPECIES OF ASPIDOSIPHON (SIPUNCULA)
FROM THE WESTERN ATLANTIC OCEAN

Edward B. Cutler

Abstract.—A new species of Sipuncula, Aspidosiphon (Aspidosiphon)
gosnoldi, is described. This form had earlier been identified erroneously as
Aspidosiphon spinalis by Cutler (1973) and Cutler and Cutler (1979). It has
been collected off the southeastern U.S., Bahamas, and Brazil at depths
between 7 and 185 m.

In Cutler (1973), and Cutler and Cutler (1979) the name Aspidosiphon
spinalis Ikeda (1904) was used for two populations of western Atlantic
Ocean worms. It was noted (Cutler, 1973:175), ‘‘This identification is made
with some reservations because no specimens of Ikeda’s material are avail-
able for comparison and his illustrations are not very clear.’’ After spending
several months in Japan examining that fauna and reevaluating the species
described by Ikeda and Sato (Cutler and Cutler, 1981) it became clear to
me that I had misused that name; 1.e., the animals from the Atlantic Ocean
were different from those Ikeda looked at. Aspidosiphon spinalis is now
considered to be a junior synonym of A. elegans. Much of the confusion
centered around the meaning of the term “‘spine.’’ This is discussed in more
detail in Cutler and Cutler (1981). The word spine has been used for two
different kinds of structure: (A) large, conical, curved in side view, dark
brown structures; (B) small, unpigmented, gently curved, single-pointed
hooks. We have restricted our usage of spine to type A, such as those
present in A. elegans, and call type B unidentate hooks, such as those
present in this species. Consultation with Mary Rice (Smithsonian Institu-
tion) and examination of the existing literature has led me to conclude that
these animals represent a new species.

Aspidosiphon (Aspidosiphon) gosnoldi, new species
Figs. 1—4

Aspidosiphon spinalis Ikeda.—Cutler, 1973:175-176, fig. 48—49.—Cutler
and Cutler, 1979:107.

Description.—As almost all of these worms are coiled, having lived in
gastropod shells, it is difficult to obtain accurate measurements (Fig. 1).
They are commonly from 5—15 mm long but range from approximately 2—22
mm. The color is cream or pale tan with shields that vary from light to dark

446 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-3. Aspidosiphon gosnoldi: 1, External view, introvert incomplete (from Cutler,
1973); 2, Unidentate and bidentate hooks from the introvert (scale 10 um); 3, Anterior end of
trunk, showing anal shield (a, anus; 1, introvert).

brown. In larger animals, the anterior third of the trunk appears brownish
because of the development of darker cuticular plaques around the papillae.
A few worms have commensal endoprocts, and others have commensal poly-
chaetes living with them in the shells.

The introvert is 2—3 times the trunk length, and its diameter decreases
near the end just behind the tentacular crown which is composed of 8—10
small, finger-shaped tentacles. No introvert was completely extended so no
comment can be made about tentacle position. The distal 10-20% bears 25-
60 complete rings of hooks followed by a short transition zone of incomplete
rings, and eventually the hooks are scattered over most of the remainder of
the introvert. The hooks near the distal end have secondary points while
the remainder are unidentate (Fig. 2) and both are about 22—26 um tall. The
introvert also carries cylindrical papillae (15-17 ym tall) between the hook
rings.

The anal shield is composed of large, brown, close-set, polygonal, ran-
domly arranged, chitinous subunits (Fig. 3). In general, it has well-defined

VOLUME 94, NUMBER 2 447

Table t.—A comparison of the known distribution of 3 similar Aspidosiphon.

A. gosnoldi A. misakiensis A. gerouldi
Ocean W. Atlantic W. Pacific E. and W. Atlantic
Latitudes 24-35°N 25—35°N 8°S-16°N
and 23°S
Common depth 10-100 m 0-5 m 5-30 m

limits. The caudal shield, composed of scattered chitinous papillae is gen-
erally paler and less well defined. These papillae give a suggestion of a radial
arrangement.

Internally the longitudinal musculature is continuous except that a few
larger worms show a suggestion of separate bundles, particularly just behind
the anal region. The 2 retractor muscles originate from the posterior end of
the body at the border of the caudal shield and merge to form one muscle
before inserting in the anterior end. These 2 muscles remain separate for
about 35-50% of the total length. The coiled intestine is anchored at both
ends by the spindle muscle but fixing muscles were not seen. The anterior
attachment of the spindle muscle is on the body wall just anterior to the
anus. The pale nephridia are about half the length of the body and are fixed
to the body wall by connective tissue for much of their length.

Relationships.—This species is in the group of Aspidosiphon (Aspidosi-
phon) which have both bidentate and unidentate hooks, 2 retractor muscles
Originating near or on the posterior end of the trunk, an ungrooved anal
shield, and lack dark, conical, spinelike papillae on the base of the introvert.
This group now includes only A. gerouldi and A. misakiensis. A. gosnoldi
differs from A. gerouldi in that its anal shield is much less compact, made
up of units which are spread apart, while A. gerouldi has one which is
compact and granular with the subunits fused. The caudal shield of A.
gerouldi is also more distinct and grooved. The difference from A. misa-
kiensis also lies in the nature of the anal shield (A. misakiensis being again
more compact and coarse). The longitudinal muscle layer of A. misakiensis
very often exhibits fractures or splits in it other than just in the anal region.
These three species are geographically separate populations with some dif-
ferences in habitat preference (Table 1). When one looks at representatives
of these three populations, they do look different but it becomes difficult to
articulate those differences precisely. It is possible that additional analyses
with more sophisticated techniques will show these forms to be conspecific.

The name of this new species acknowledges the vessel used by the U.S.
Geological Survey to collect most of these animals, the R/V Gosnold. The
holotype is from Sta. 1692, 12 June 1964, 29°30'N, 80°29’W, 32 m, and is
deposited at National Museum of Natural History, Smithsonian Institution,
USNM 61624. Paratypes are from R/V Eastward St. 1439, collected by I.

448 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Recorded occurrences of Aspidosiphon gosnoldi off the southeastern United States.
For records in the Bahamas and Brazil, see text.

VOLUME 94, NUMBER 2 449

E. Gray and L. McCloskey, 21 May 1965, 34°59’N, 75°28’ W, 100 m, USNM
61625.

Distribution.—In the Northern Hemisphere (Cutler, 1973, 287 specimens
from 103 stations) this species inhabits the continental shelf from Cape Hat-
teras to the Florida Keys (Fig. 4). The known latitudinal range is from 24°27’
to 35°00’N. Seventy-one percent of the records are from depths less than
35 m, 17% between 35-85 m, and 12% greater than 85 m. The known depth
range is 7-185 m; the temperatures range from 15—28°C. Most of these
animals live in empty mollusk shells. One previously unpublished record is
of 2 worms collected by B. Thomassin from the Bahamas in 1974, seaward
reef slope in sand at 15 m. The southern population (Cutler and Cutler,
1979) is known from 2 stations off Brazil (23°S) at 103 m (11 Specimens) and
25 m (1 specimen) collected by the Calypso.

Literature Cited

Cutler, E. B. 1973. Sipuncula of the Western North Atlantic.—Bull. American Mus. Nat.
Hist. 152:105-204.

, and N. J. Cutler. 1979. Campagnes de la Calypso au large des cotes Atlantiques
Africaines (1956 et 1959) et Sud-Americaines (1960-1961). 23. Sipuncula.—Resultats
Sci. Camp. de la Calypso 11:103-109.

, and - 1981. A reconsideration of Sipuncula named by I. Ikeda and H. Sato.—
Publ. Seto Mar. Biol. Lab. 26:51-93.

Ikeda, I. 1904. The Gephyrea of Japan.—Jour. Coll. Sci., Imp. Univ. Tokyo 20: 1-87.

Utica College of Syracuse University, Utica, New York 13502.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 450-454

PSOLIDIUM PROSTRATUM, NEW SPECIES, FROM
OFF THE EAST COAST OF THE U.S.A.
(ECHINODERMATA: HOLOTHUROIDEA)

David L. Pawson and John F. Valentine

Abstract.—A new species of Psolidium is described. It was collected off
the North Carolina and South Carolina coasts at depths ranging from 383
to 450 meters. This is the first record of the genus Psolidium from the
Western North Atlantic.

The genus Psolidium Ludwig, 1886, as presently characterized includes
approximately 25 species. Psolidium is widely distributed in the polar re-
gions and the Pacific Ocean, to depths of approximately 3000 meters. Until
now, only a single species, P. arcuatum (Herouard, 1912), from off the
Azores, was known from the Atlantic Ocean. The present new species is,
therefore, the first record of the genus from the Western North Atlantic.

As psolids usually attach themselves to hard substrata, such as rocks, it
is Obvious that they would be more plentiful in rocky habitats that are
usually avoided by research vessels engaged in trawling and dredging. It is
thus likely that several species of psolids, and especially Psolidium, remain
to be discovered in rocky habitats in the Western North Atlantic.

The new species was collected off South Carolina by Texas Instruments
Ecological Services during 1977, while undertaking the South Atlantic
Benchmark Program for the Bureau of Land Management. An additional
specimen collected from off North Carolina was found in the reference
collection of the Duke University Marine Laboratory, Beaufort, North Car-
olina.

Family Psolidae Perrier, 1902
Psolidium Ludwig, 1886
Type-species.—Psolidium dorsipes Ludwig, 1886.

Psolidium prostratum, new species
Figs. 1, 2

Diagnosis.—Small species, probably not exceeding 20 mm in total length.
Numerous tube feet scattered on dorsal surface. Oral and anal apertures sur-

VOLUME 94, NUMBER 2 451

eee Ss

IE

o, @es.

yr SJ
seta Reyes

Fig. 1. Psolidium prostratum: Holotype, dorsal aspect. Drawing by C. G. Messing.

rounded by numerous scales; no identifiable oral or anal valves. Approxi-
mately 12-20 scales between oral and anal apertures. Mid-ventral radius
with few (4—5) tube feet. Ossicles in sole exclusively knobbed plates, usually
with more than 4 perforations; no cups.

Material examined.—10 specimens and 5 fragments from the following
localities: HOLOTYPE.—USNM _~ E20993, 32°20'06"N, 78°11'04’”W,
412 meters, 15 February 1977, temperature 17.5°C, salinity 35.2%, dissolved
O, 4.9 mg/liter (Sta. 159-2), total length 11.5 mm. PARATYPES.— USNM
E20994, same data as Holotype, 2 specimens, 10.5, 3.5 mm; USNM E20995,
32°19'59"N, 78°10'39.06"W, 383 meters, 12 May 1977, temperature
8.79°C, salinity 35.3%o, dissolved O, 4.7 mg/liter, 6 specimens and 5 frag-
ments, total length 2-13 mm (Sta. 501-2 to 501-6). Other material: Duke
University Marine Laboratory, 34°17.5’N, 75°49.5’W, 450 meters, 13 March
1965, 1 specimen, total length 6 mm.

Description.—Body very flattened, contorted to conform to shape of sub-
strate. Total length 1.5-15 mm. Width approximately 50% of length. Dorsal
surface with numerous thin, fragile scales; at least 12-20 scales between
oral and anal apertures. Scales smooth, approximately equal-sized, aver-
aging 1 mm in diameter. Scales decrease sharply in size towards ambitus.
Tube feet scattered over dorsal surface, at least 100 feet in large specimens;
feet emerge between and through dorsal scales (Fig. 2C). Oral and anal
apertures similar, oral aperture larger. Both surrounded by numerous scales

452 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Psolidium prostratum, ossicles: A, Knobbed buttons from sole; B, Perforated plates
from tentacles; C, Outline drawings of dorsal plates, showing holes for passage of tube feet;
D, Endplate from ventral tube foot; E, Plates from dorsal tube feet; F, Developing buttons
from sole of 1.5 mm total length juvenile; G, Endplate from tube foot of 1.5 mm total length
juvenile. The scale on Fig. 2A also applies to figs. 2B, D-G.

VOLUME 94, NUMBER 2 453

without regular arrangement. Anal aperture can be pushed outwards on a
small anal cone. Boundary between dorsal and ventral surfaces marked by
a sharp edge. Sole fragile, transparent, with scattered marginal tube feet.
Midventral radius also with scattered tube feet, usually only 4-5 present.
Tentacles weakly dendritic in larger specimens. Smaller specimens of less
than 12 mm length often with unbranched tentacles; in these, primordial
branches often evident. Tentacles 10, white, 2 ventral tentacles smaller than
others. Color of body usually white, although in few specimens dorsal scales
are light to dark brown.

Ossicles of sole exclusively knobbed buttons of average length 118 pm;
usually there are more than 4 perforations (Fig. 2A). In sole of 1.5 mm long
juvenile, ossicles smooth, with few perforations (Fig. 2F). Ventral tube feet
with well developed endplates (Fig. 2D), even in juveniles (Fig. 2G). Dorsal
feet lack endplates, but contain some curved simple plates, sometimes with
a few small knobs (Fig. 2E). Tentacles contain small perforated plates and
rods, often curved, of variable size, ranging from approximately 75 yum long
to approximately 400 um long (Fig. 2B).

Ecology.—Specimens were recovered from box core samples, in which
they were found attached to pieces of volcanic agglomerate estimated to be
between 1 million and 10 million years old. Other faunal elements included
Tharyx sp., Thelepus cincinnatus, and several species of amphipods.

Relationships.—The only other known Atlantic congener, P. arcuatum
Herouard, has smooth plates in the sole with few perforations. From the
eastern Pacific, in the vicinity of Central America, 6 Species are known.
They differ from P. prostratum in the following ways: P. gracile Ludwig,
1894, has sole plates with fewer perforations and many fewer knobs. P.
panamense Ludwig, 1894, has a higher, almost cylindrical body, much less
conspicuous dorsal feet, and much larger sole plates. P. ekmani Deichmann,
1941, has pearl-like projections dorsally and 4-holed ossicles in the sole. P.
eubullatum Deichmann, 1941, has heavy buttons and plates ventrally, and
the dorsal feet have no ossicles. P. dorsipes Ludwig, 1886 and P. planum
Deichmann, 1941, both have cups in addition to the other ossicles in the
sole. The present new species appears to bear no close relationship with
other known species in the genus.

Acknowledgments
We wish to thank Mr. J. Kevin Shaw and Ms. Debbie L. Blizzard for
their help in many ways.

Literature Cited

Deichmann, E. 1941. The Holothurioidea collected by the Velero III during the years 1932
to 1938. Part I, Dendrochirota.—Allan Hancock Pacific Expeditions 8(3):61-194, pls.
10-30.

454 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Herouard, E. 1912. Holothuries nouvelles des campagnes du yacht Princesse-Alice.—Bulletin
de l'Institut Océanographique 239:1-9, figs. 1-7.

Ludwig, H. 1886. Die von G. Cheirchia auf der Fahrt der Kgl. Ital. Corvette “‘Vettor Pisani’’
gesammelten Holothurien.—Zoologische Jahrbucher 2:1—36, pls. 1-2.

.1894. The Holothurioidea.—Memoirs of the Museum of Comparative Zoology 17(3):1-
183, pls. 1-19.

(DLP) Department of Invertebrate Zoology, National Museum of Natural
History, Smithsonian Institution, Washington, D.C. 20560 U.S.A.; (JFV)

Barry Vittor & Associates, 8100 Cottage Hill Rd., Mobile, Alabama 36609,
U.S.A.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 455-457

A NEW SPECIES OF HOMOCHAETA (OLIGOCHAETA:
NAIDIDAE) FROM THE WEST COAST OF CANADA

Peter M. Chapman

Abstract.—Homochaeta raptisae n. sp. is described from the lower Fra-
ser River, British Columbia. The new species represents the first confirmation
of the genus Homochaeta in North America.

The genus Homochaeta is distinguishable from other naidid genera by
the fact that its dorsal setae start from segment II and hair setae are absent.
The genus presently contains two recognized species identified from Eurasia
and a third species, H. lactea, described from a single specimen collected
in South America (Cernosvitov, 1937) and considered to be species incerta
sedis (Brinkhurst and Jamieson, 1971). This paper describes a new species
of Homochaeta from the North American west coast, Homochaeta rapti-
sae.

Materials and Methods

Specimens of Homochaeta raptisae were identified from oligochaetes col-
lected at several stations in the Tilbury Slough area of the lower Fraser
River, B.C. (Chapman, 1980) during a baseline study by staff at the Habitat
Protection Division of the Department of Fisheries and Oceans Canada
(principal investigator, Mr. M. Nassichuk). Preserved worms were stained
in paracarmine, and mounted whole in Canada balsam. The holotype and
paratypes are deposited at the British Museum (Natural History) (BMHN).
Additional material is deposited at the United States National Museum of
Natural History (NMNH), and the National Museums of Canada (NMCIC).

Homochaeta raptisae, new species
Fig. 1

Material.—Holotype: BMNH 1980.47.1, whole mounted specimen. Para-
types: BMNH 1980.47.2-3, 2 specimens in alcohol. NMCIC 1980-1527, 2
specimens in alcohol; 1980-1528, whole mounted specimen. NMNH 062010,
2 specimens in alcohol; 062011, whole mounted specimen. Additional ma-
terial: In the collection of P. M. Chapman, 30 specimens from various lo-
cations in the lower Fraser River, B.C.

Type-locality.—Lower Fraser River, B.C., Canada; subtidal silty sedi-
ment; 49°8.9’N, 122°59.7’W, collected 26 April 1977.

Etymology.—Named for my wife, Stavroula Raptis.

Description.—No pigment or eyes. Up to 15 segments, length (preserved)
0.5-1.0 mm. Prostomium elongate, triangular and upturned. All setae of

456 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

0.017

Fig. 1. Homochaeta raptisae: A, Outline of whole-mounted worm; B, Dorsal anterior setae;
C, Ventral setae. All lengths in mm.

same shape, teeth of equal length, upper tooth thinner than lower, nodulus
distal; dorsal anterior teeth with a single intermediate pectination; 5 setae
per bundle in II, in other segments 3. Pharynx in II-III, stomach in V.
Chloragogen from V. Coelomocytes absent. Evidence of budding.

Discussion

The genus Homochaeta is so distinctive, with bifid dorsals starting from
II and lacking hair setae, that it cannot be confused with other naidids. With
the discovery of H. Raptisae n. sp., the genus is now expanded to include
four species, with the Eurasian forms (H. naidina and H. setosa) geograph-
ically separated from the South American species (H. lactea) and the pres-
ent new species. A report of H. naidina from Virginia (Falls 1974) is un-
confirmed, as voucher specimens were lost (Hiltunen and Klemm 1980).

H. raptisae can be distinguished from the other three species in the genus
based on size, prostomial shape, and setal shape and number. Examination
of the unique holotype of H. lactea (sent to the author courtesy of Mr. R.
W. Sims of the British Museum) confirmed that H. raptisae is a new species.
The other two species in the genus are sufficiently distinct that an examina-
tion of type-specimens was not necessary.

VOLUME 94, NUMBER 2 457

All specimens of H. raptisae examined contained large numbers of dia-
toms in their guts, but very little sediment, which suggests that they were
feeding selectively on periphyton. All specimens were immature, a not un-
usual observation among the Naididae, which reproduce asexually by bud-
ding. Thus a description of sexually mature specimens must await the col-
lection of further material.

H. raptisae was collected at the type-locality in numbers varying from
<500 m™ in winter and summer to >10,000 m~® in spring. This seasonal
abundance pattern, typical of many naidids, will be discussed elsewhere and
is documented by Chapman (1980). The relatively large numbers of H. rap-
tisae observed in the Fraser River suggest that this species will be identified
from other areas in future. Its absence from previous taxonomic records
may be due partly to the fact that inexperienced taxonomists could mistake
the species for an immature tubificid, since its dorsals start on II. However,
it is more probable that, due to its small size coupled with the common use
of 0.5 mm sieves in benthic collections, previous workers have failed to
collect or sort H. raptisae from sites where it might be abundant. The worms
described in the present study were sorted using a 0.250 mm sieve; however,
oligochaete collections made in 1972 and 1973 from the type-area (Westwa-
ter station 6—Northcote et al., 1976), sorted with a 0.5 mm sieve and ex-
amined by the author, did not include H. raptisae. This omission serves to
emphasize the importance of sieve size in base-line benthic collections.

Acknowledgments

I am indebted to Dr. R. O. Brinkhurst for his assistance with the taxo-
nomic descriptions, and I thank Ms. B. Piercey and Mr. M. Nassichuk who
collected and made available the oligochaete material used in this study.

Literature Cited

Brinkhurst, R. O., and B. G. M. Jamieson. 1971. Aquatic Oligochaeta of the World.—U. of
Toronto Press, Toronto. 860 pp.

Cernositov, L. 1937. Notes sur les Oligochaeta (Naididées et Enchytraeidées) de |’ Argen-
tine.—An. Museo Nac. Buenas Aires 39: 135-157.

Chapman, P. M. 1980. Identification and analysis of oligochaete samples collected from the
lower Fraser River, Feb.—Aug. 1977.—Report prepared for Habitat Protection, Fisheries
and Oceans Canada. 13 pp.+ appendices.

Falls, E. Q. 1974. A taxonomic survey of freshwater oligochaetes from the Richmond Vir-
ginia area with reference to commensal ciliates.—Va. J. Sci. 25:25-29.

Hiltunen, J. K., and D. S. Klemm. 1980. A guide to the Naididae (Annelida: Clitellata:
Oligochaeta) of North America.—EPA-600/4-80-031. 48 pp.

Northcote, T. G., N. T. Johnston, and K. Tsumura. 1976. Benthic, epibenthic and drift fauna
of the lower Fraser River.—Westwater Technical Report 11, U. of British Columbia.
227 pp.

E.V.S. Consultants Ltd., 195 Pemberton Avenue, North Vancouver,
B.C., V7P 2R4 Canada.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 458-461

FIRST DESCRIPTION OF THE MALE OPPOSUM SHRIMP,
HETEROM YSIS BERMUDENSIS BERMUDENSIS
(CRUSTACEA: MYSIDACEA)

Thomas E. Bowman

Abstract. —Some characters of Heteromysis bermudensis bermudensis
are described and illustrated, from Bermudan specimens. Differences in the
pleopod 4 support Bacescu’s (1968) recognition of a subspecies, H. b. ce-
sari, for Cuban specimens.

The opossum shrimp, Heteromysis bermudensis Sars, 1885, has been
referred to by 6 different authors, but because of scanty material, it remains
inadequately characterized. In particular, the ¢ of the typical subspecies is
unknown. Sars (1885) had only a single 2 with all the legs broken off. Verrill
(1923) had an unspecified number of specimens, presumably collected by
his students and him in 1898 and 1901, since the color in life was noted.
Tattersall (1951) had 2 ‘“‘rather badly damaged’’ 2 2 (an understatement!),
USNM 82388, collected in 1876-77 by G. Brown Goode. Clarke (1955) also
examined these damaged 2 @ in addition to unspecified *‘fresh material.”’
Bacéescu (1968) had 4 66 and Brattegard (1973) 1 @.

The Sars, Verrill, and Clarke specimens came from Bermuda. Bacescu’s
66, from Cuba, were assigned to a new subspecies, H. b. cesari, recently
reported also from the Saba Bank by Brattegard (1980). Brattegard’s (1973)
?, from Colombia, was identified as H. bermudensis, and presumably be-
longs to the typical subspecies. The latter was called H. b. typica by Ba-
cescu, but in accordance with ICZN Art. 47a the name should be dH. b.
bermudensis.

For the present study I have had available 41 ¢ 2 from Hamilton Harbor,
Bermuda, washed from algae and a sponge collected at depths of 2—6 feet
by Lanelle W. Peterson, 13 August 1961.

Because the ¢ of H. b. bermudensis was unknown, Bacescu was unable
to employ the useful ¢ secondary sexual characters in distinguishing the
subspecies. In Heteromysis, as discussed in detail by Tattersall (1967), pleo-
pods 2, 3, and especially 4 may be modified in the 6, thereby providing
valuable taxonomic characters. The characters used by Bacescu in a table
comparing the subspecies are size, number of spines on the margin and in
the sinus of the telson, color, length of the scale of antenna 2, and shape of
eyes. A discussion of these follows:

Size: H. b. cesari is smaller (total length 4.0-4.2 mm) than H. b. ber-

VOLUME 94, NUMBER 2 459

mudensis. Sars’ holotype measured 6 mm; Verrill and Clarke reported 6-8
and 7-9 mm respectively for their specimens. However, Brattegard’s 2 was
only 4.6 mm, and the Peterson specimens are 4.5—4.8 mm, not much larger
than H. b. cesari.

Marginal spines of telson: Including the 2 apical spines, H. b. bermuden-
sis has 8 (Sars, Brattegard), 7—9 (Clarke), 8—9 (Peterson specimens); H. b.
cesari has 8 (Bacescu listed 6, but did not include the 2 apical spines).

Spines in sinus of telson: H. b. bermudensis has 18 (Sars), 18—20 (Clarke),
16 (Brattegard), 15S—16 (Peterson specimens); H. b. cesari has 14 (figures
given in Bacescu’s table are reversed).

Length of scale of antenna 2: According to Bacescu, in H. b. bermudensis
the scale extends beyond the endopod peduncle and in H. b. cesari the
scale is shorter than the peduncle. But Sars described the scale as ‘‘ex-
ceedingly small, scarcely as long as the antennular peduncle,’’ I believe
‘‘antennular’’ is a lapsus for ‘‘antennal,’’ for in Sars’ pl. 38, figs. 1 and 2,
the scale just reaches the end of the antennal peduncle, but is much shorter
than the antennular peduncle. The apparent lengths of the antennal peduncle
and scale depend to some extent upon the angle at which they are viewed
and upon whether or not there is distortion from cover glass pressure.

Sars noted a distal segment in the antennal scale. Bacescu showed a distal
segment in his figure of the head region but not in his more enlarged figure
of antenna 2, and remarked, **. . . on ne distingue pas clairment d’article
apical.’ Brattegard found no distal segment in his Colombian specimens,
but it is clearly present in the Peterson specimens (Fig. Ic).

Eyes: Bacescu referred to the eyes of H. b. bermudensis as more or less
cylindrical and to those of H. b. cesari as irregularly globular. This distinc-
tion is not clearly evident from a comparison of the figures of Sars, Bacescu,
and Brattegard, but other differences are apparent. In H. b. cesari the
cornea occupies nearly half of the eye, in H. b. bermudensis only 4. Also,
the small concavity in the proximolateral part of the cornea in H. b. cesari
has not been observed in H. b. bermudensis.

The above discussion suggests that most of the criteria used by Bacescu
to distinguish the 2 subspecies either overlap or differ slightly, and hence
are of questionable reliability. Because the 6 of H. b. bermudensis was
unknown, Bacescu was unable to use ¢ secondary sexual characters. How-
ever, we can now add to Bacescu’s criteria the anatomy of the d pleopod
4, which differs in the 2 subspecies as follows:

Spines/setae H. b. bermudensis H. b. cesari
Lateral margin yy 0
Medial margin, proximal to

pseudobranchial lobe 1 0
Anterior surface 6 2

Distal margin ca 35, with flagellum 26-30 without flagellum

460 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

NYY
Wael
WS

Fig. 1. Heteromysis bermudensis bermudensis: a, 6 Antenna 1, sympod, dorsal; b, Distal
end of antenna | sympod, ventral, showing 3 lobe; c, ¢ antenna 2; d, Thoracic endopod 3,
distal segments; e, Thoracic endopod 4; f, Pleopod 3, 3; g, Pleopod 4, ¢; h, Telson.

VOLUME 94, NUMBER 2 461

These differences, together with those of the eyes, seem sufficient to
support recognition of the 2 subspecies. Indeed, Ba¢escu remarked that if
differences in the ¢ pleopod 4 should be found, the Cuban form would merit
recognition as a full species. I quite agree, but believe this step should be
deferred until pleopod 4 of 6¢ of other populations can be examined for
variability.

In other respects the Peterson specimens agree well with the Colombian
© described by Brattegard except for the presence of a distal segment in
the scale of antenna 2. The ¢ lobe of antenna | is very short and densely
covered with fine setae (Fig. 1b). The carpopropus of thoracic endopod 4
is 3-4 segmented (Fig. le); those of endopods 5-8 are 5—7-segmented. Sex-
ual dimorphism of the pleopods is found only in pleopod 4 (Fig. 1g).

Acknowledgments

I am grateful to Austin B. Williams for reviewing the manuscript.

Literature Cited

Bacescu, Mihai. 1968. Heteromysini nouveaux des eaux Cubaines: Trois especes nouvelles
de Heteromysis et Heteromysoides spongicola n.g. n. sp.—Revue Roumaine de Biol-
ogie, seria Zoologie (13)4:221-—237.

Brattegard, Torleiv. 1973. Mysidacea from shallow water on the Caribbean coast of Colom-

bia.—Sarsia 54: 1-66.

. 1980. Platymysis facilis gen. et sp. nov. (Crustacea: Mysidacea: Heteromysini) from

the Saba Bank, Caribbean Sea.—Sarsia 65:49—52.

Clarke, William D. 1955. A new species of the genus Heteromysis (Crustacea, Mysidacea)
from the Bahama Islands, commensal with a sea-anemone.—American Museum Novi-
tates, No. 1716:1-13.

Sars, G. O. 1885. Report on the Schizopoda collected by H. M. S. Challenger during the
years 1873—76.—Report on the Scientific Results of the Voyage of H. M. S. Challenger
during the Years 1873-76, Zoology 13(37):1—228, pls. 1-38.

Tattersall, Olive S. 1967. A survey of the genus Heteromysis (Crustacea: Mysidacea) with
descriptions of five new species from tropical coastal waters of the Pacific and Indian
Oceans, with a key for the identification of the known species of the genus.—Trans-
actions of the Zoological Society of London 31:157-193.

Tattersall, Walter M. 1951. A review of the Mysidacea of the United States National Mu-
seum.—United States National Museum Bulletin 201:I-X + 1-292.

Verrill, A. E. 1923. Crustacea of Bermuda. Schizopoda, Cumacea, Stomatopoda and Phy]l-
locarida.—Transactions of the Connecticut Academy of Arts and Sciences 26:181-211,
pls. 49-52.

Division of Crustacea, Smithsonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 462-463

COMMENTS ON THE TAXONOMIC STATUS OF
TOMMOTIA MISSARZHEVSKY 1970
(TOMMOTIDAE: MICROPROBLEMATICA)

Richard W. Huddleston

Missarzhevsky in Rozanov and Missarzhevsky (1966) established the gen-
era Camena and Camenella for problematic microfossils of lower Cambrian
age. The generic name Camena Missarzhevsky in Rozanov and Missar-
zhevsky 1966 was preoccupied by Camena Hewitson 1865 and subsequently
Missarzhevsky (1970) proposed the replacement name Tommotia for the
junior homonym. Camena Missarzhevsky in Rozanov and Missarzhevsky
had been made the type-genus of the family Camenidae by Missarzhevsky
in Rozanov et al. (1969).

Bengtson (1970) pointed out that Tommotia and Camenella were con-
generic, representing separate parts of the same organism. Bengtson (1970)
corrected the family name from Camenidae to Tommotiidae based on the
new name Tommotia, citing authorship to Missarzhevsky 1970. Bengtson
(1970:369-370) commented, ‘‘Since Tommotia is the replacement name for
the type-genus of a family, it will—contrary to what is normally the case—
take the date of the name replaced, Camena (article 39 of the International
Code of Zoological Nomenclature), and thus be able to compete in priority
with Camenella, . . . Accordingly, the generic name Tommotia, because of
its status as type-genus for the family, is here selected as the valid name for
the combined genera.’ However, ICZN article 39 only indicates that the
family name Camenidae is invalid because of the junior homonym status of
its type-genus, Camena. Further, ICZN article 60(b) indicates that Tom-
motia must compete with Camenella with its 1970 date rather than the date
of the name it replaced, 1966. Regardless of the fact that Tommotia is the
replacement name for the type-genus it is a junior subjective synonym of
Camenella and must be treated as such by all who recognize the synonymy.
(ICZN article 60).

Acknowledgments

I thank D. Haman, Chevron Oil Field Research Company for comments
and review of the manuscript, and Chevron Oil Field Research Company
for permission to publish.

VOLUME 94, NUMBER 2 463

Literature Cited

Bengtson, S. 1970. The lower Cambrian fossil Tommotia.—Lethaia 3:363-392.

Missarzhevsky, V. V. 1970. Renaming of an early Cambrian genus of phosphatic problematic
organisms.—Paleont. Zhur. 1970(4):100.

Rozanov, A. Yu., and V. V. Missarzhevsky. 1966. Biostratigraphy and fauna of the oldest
Cambrian horizons.—Akad. Nauk SSR. Geol. Inst. Trudy, 148, 126 pp.

Rozanov, A. Yu. et al. 1969. The Tommotian stage and the Cambrian lower boundary prob-
lem.—Akad. Nauk. SSR. Geol. Inst. Trudy 206, 380 pp.

Chevron Oil Field Research Company, P.O. Box 446, La Habra, Califor-
nia 90631.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 464-478

NOTES ON WEST AFRICAN PIPEFISHES
(SYNGNATHIDAE), WITH DESCRIPTION
OF ENNEACAMPUS, N. GEN.

C. E. Dawson

Abstract.—Syngnathus pellegrini Fowler, described from Gabon, is based
on mislabeled specimens from the western north Atlantic and is a junior
synonym of S. fuscus Storer. The name Enneacampus is proposed for a
new syngnathine (tail-pouch) genus characterized, in part, by a 9-ray caudal
fin and by the presence of an anal fin, pouch-protective plates, and everted
pouch-closure. The type-species (Syngnathus ansorgii Boulenger 1910) and
its only congener, Enneacampus kaupi (Bleeker) are rediagnosed and illus-
trated; Syngnathus pulchellus Boulenger and S. olssoni Johnels are junior
synonyms of Enneacampus ansorgii. Oostethus brachyurus aculeatus
(Kaup) and the two species of Enneacampus are the only pipefishes known
to occur commonly in shallow coastal or inland waters of west Africa be-
tween Sénégal and Angola (ca. 16°N—18°S); all are known to breed in fresh
water.

The genus Syngnathus Linnaeus (type-species S. acus L.) has long been
a catchall for pipefishes of varying lineages or uncertain taxonomic status.
The number of nominal species is variously estimated at 40-50 or more, but
the number of valid species more closely approaches 30. Studies on western
African pipefishes currently referred to Syngnathus show that one name is
a junior synonym of an extralimital species and that four nominal species,
apparently freshwater forms, are not referrable to Syngnathus s.s. I here
clarify the identity of S. pellegrini Fowler, propose a new genus for the
accommodation of S. ansorgii Boulenger and S. kaupi Bleeker, and refer
two nominal species (S. pulchellus and S. olssoni) to the synonymy of S.
ansorgil.

In addition to species rediagnosed here (ansorgii and kaupi), preliminary
studies indicate that only one other pipefish [Oostethus brachyurus aculea-
tus (see Dawson, 1979)] occurs commonly in the shallow coastal and inland
waters of western Africa between Sénégal and Angola (ca. 16°N—18°S).
Compared to the western Atlantic, with 14 coastal and inland species known
from the same latitudes, western Africa supports a depauperate pipefish
fauna.

VOLUME 94, NUMBER 2 465

Methods and Materials

Measurements (mm) are of total length (TL), standard length (SL) or head
length (HL). Unless otherwise noted, color descriptions are from specimens
preserved in alcohol; the term ‘‘venter’’ refers to the ventral surface of the
head or body; other methods follow Dawson (1977). Abbreviations for re-
positories of material examined are: AMNH, American Museum of Natural
History; ANSP, Academy of Natural Sciences, Philadelphia; BMNH, Brit-
ish Museum (Natural History), London; CAS-SU, Stanford University col-
lections, how housed at California Academy of Sciences; GCRL, Gulf Coast
Research Laboratory Museum; IRSNB, Institut Royal des Sciences Natu-
relles de Belgique, Brussels; MCZ, Museum of Comparative Zoology, Har-
vard University; MNHN, Muséum National d’Histoire Naturelle, Paris;
NRM, Naturhistoriska Riksmuseet, Stockholm; RMNH, Rijksmuseum van
Natuurliyke Historie, Leiden; UMMZ, Museum of Zoology, University of
Michigan; USNM, National Museum of Natural History, Smithsonian In-
stitution; ZMUC, Zoological Museum, University of Copenhagen.

Syngnathus pellegrini Fowler

Fowler (1919) described this species from four specimens (88-113 mm
TL), reportedly collected in Gabon by P. B. DuChaillu. The name was listed
by Fowler (1922) and he later (1936) gave the type locality as “‘coast of the
French Congo.’’ Additional specimens referrable to S. pellegrini have not
been taken along the African coast. The validity of this species was ques-
tioned by Blache (1962) but it was included in a recent key to eastern At-
lantic fishes (Blache et al., 1970).

The original description gives counts of 18 + 38 rings, 38 dorsal-fin rays
and 4 + 6 subdorsal rings for the holotype (ANSP 975) and 18-20 + 37-38
rings and 35-37 dorsal-fin rays for the three paratypes (ANSP 976-978). My
counts from these juveniles (now 83-107 mm SL) follow: rings 19 + 37-38;
dorsal-fin rays 36, 37, 41, 42; pectoral-fin rays 13 (6 counts), 14 (2); caudal-
fin rays 10; subdorsal rings 5-4 + 4.5-5.5. The configuration of principal
head and body ridges is that of the genus Syngnathus, and the anal fin is
present. Pertinent proportional values are: HL in SL 8.1-8.4, snout length
in HL 2.1—2.2, snout depth in snout length 4.5 (in holotype), length of dorsal-
fin base in HL 0.8-0.9.

These specimens are conspecific with S. fuscus Storer, a common species
along the Atlantic coast of the United States. There have been a number of
labeling problems with material thought to belong to the DuChaillu (or Du-
chaillu) west African collection and mislabeling of these western Atlantic
pipefishes is not surprising. Fowler (1936) noted that Ischnomembras ga-
bunensis Fowler 1903 is conspecific with the American atherinid Menidia

466 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

menidia (Linnaeus), and I am advised (in litt.) by Mrs. Eugenia Bohlke
(ANSP) that at least three other species, treated as west African by Fowler
(1919), are based on misidentified specimens of western Atlantic origin.

Enneacampus, new genus

Type-species.—Syngnathus ansorgii Boulenger 1910.

Diagnosis.—Superior trunk and tail ridges discontinuous near rear of dor-
sal fin, inferior trunk and tail ridges continuous, lateral trunk and tail ridges
discontinuous below dorsal-fin base; venter of trunk clearly V-shaped, the
median longitudinal ridge with or without a prominent fleshy keel. Snout
length 1.8-—2.8 in HL; median dorsal snout ridge not strongly elevated, usu-
ally ending on interorbital but sometimes continued to frontal ridge, its
margin entire to finely denticulate in subadults and adults; interorbital de-
pressed mesially but flared dorsolaterad toward orbits; median dorsal head
ridges a little elevated; supraopercular ridge usually distinct; opercle with
a complete, straight, longitudinal ridge, otherwise smooth or ornamented
with fine striae; pectoral-fin base with superior and inferior ridges. Principal
body ridges distinct but not strongly elevated, indented between rings, the
margins finely denticulate to entire; posterior angles of tail rings not pro-
duced to spine-like points; scutella without longitudinal keels. Head and
body devoid of dermal flaps, spines or prominent denticulations. Trunk rings
12-14 (mostly 13), total rings 44-50, dorsal-fin rays 22—29, pectoral-fin rays
12-17 (mostly 12-16), anal-fin rays 2-3, caudal-fin rays typically 9, the fin
rounded distally. Dorsal-fin origin on last trunk ring or first tail ring, the fin-
base not elevated, total subdorsal rings 5.0-6.5. Brood pouch below 12-18
tail rings; pouch plates present, angled a little laterad in brooding males;
pouch-closure the everted type of Herald (1959); without odontoid processes
in jaws (Dawson and Fritzsche, 1975) or bony inclusions in gill membranes
(Dawson, 1978). Maximum size at least 173 mm TL.

Comparisons.—The principal body ridge configuration of Enneacampus
(Fig. 1) is shared with several other syngnathine (tail-pouch) genera, but
Enneacampus clearly differs in having 9 rather than 10 caudal-fin rays. In
general appearance, species of Enneacampus (Figs. 2, 3) are most similar
to those of Bryx Herald, Parasyngnathus Duncker, and Syngnathus s.s.
Enneacampus also differs from Bryx in having an anal fin, and from Par-
asyngnathus in having well-developed pouch plates. Enneacampus has a
complete opercular ridge, a supraopercular ridge, and everted pouch-clo-
sure, whereas species of Syngnathus lack the supraopercular ridge, lack a
complete opercular ridge in subadults and adults, and pouch-closure is in-
verted. Furthermore, species of Enneacampus (2 specimens examined) lack
the 3rd epibranchial and have one infrapharyngobranchial, whereas the 3rd
epibranchial and two infrapharyngobranchials are present in Syngnathus
(Fritzsche 1980).

VOLUME 94, NUMBER 2 467

Fig. 1. Enneacampus ansorgii. Upper pair.—Lateral and dorsal aspects of head and an-
terior 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 110.5 mm SL holotype (BMNH 1911.6.1.129).

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

VOLUME 94, NUMBER 2 469

The Enneacampus body ridge configuration is also shared with two gas-
trophorine (trunk-pouch) genera, the monotypic Australian Leptoichthys
Kaup and the Indo-Pacific Doryichthys Kaup. Compared to the former, Ennea-
campus has fewer anal- and caudal-fin rays (respectively, 2—3 and 9 against
5 and 11) and lacks the very elongate snout and caudal fin characteristic of
Leptoichthys. Enneacampus shares the 9-ray caudal fin with Doryichthys
but differs from this trunk-pouch genus in having fewer anal-fin rays (4 rays
in Doryichthys) and in having rounded pectoral fins (usually emarginate in
Doryichthys).

Etymology.—From the Greek ennea (nine) and kampos (sea-animal), in
allusion to the characteristic 9-ray caudal fin; gender, masculine.

Remarks.—Among pipefishes which retain the caudal fin from postlarva
to adult, the number of caudal-fin rays is a highly conservative character
useful in defining genera, and the typical number of rays ranges from 8 to
11 (except when damaged or regenerated). The majority of syngnathine and
some gastrophorine genera have 10 caudal-fin rays, whereas the 9-ray fin is
less common. Nine caudal-fin rays are characteristic of several trunk-pouch
genera (e.g. Doryichthys, Oostethus Hubbs) but, among tail-pouch forms,
the 9-ray fin is found only in Enneacampus and some Indo-Pacific species
currently referred to Trachyrhamphus Kaup and Yozia Jordan and Snyder.
In Atlantic waters, the 9-ray caudal fin is found only in the two species of
Enneacampus and in eastern and western Atlantic populations of the wide-
spread doryrhamphine Oostethus brachyurus (Bleeker).

Most pipefishes with the 9-ray caudal fin (except those referred to Tra-
chyrhamphus or Yozia) are known to breed in fresh water. Although some
with a 10-ray fin may also breed in fresh water (e.g. Syngnathus scovelli
(Evermann and Kendall) and species of Pseudophallus Herald), the majority
of pipefishes with other than 9 caudal-fin rays breed in estuarine or marine
habitats.

Enneacampus ansorgii (Boulenger)
Figs. 1-2

Syngnathus ansorgii Boulenger, 1910:559 (orig. descr.; Quanza (= Cuanza)
River, Angola); Boulenger, 1915:86, fig. 73 (in key, descr.); Clausen,
1956:227 (comparisons); Daget and IItis, 1965:185 (in key); Thys, 1965:317
(listed, Fernando Poo); Thys, 1967:82, fig. 24 (descr., comparisons).

Syngnathus pulchellus Boulenger, 1915:88, fig. 74 (orig. descr., Gabon and

<_—
Fig. 2. Enneacampus ansorgii. Top to bottom: BMNH 1911.6.1.129 (110.5 mm SL, male,

holotype). UMMZ 187910 (94 mm SL, male). GCRL 15532 (110 mm SL, adult female). GCRL
15485 (ventral aspect of brood pouch of 82.5 mm SL male, illustrating disposition of pouch-
young and brood-pouch folds).

470 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

oh ahs HT FREE SW Serr Sg nd ptr ee

Fig. 3. Enneacampus kaupi. CAS-SU 63059: Top.—141 mm SL, brooding male. Bottom.—
132 mm SL, adult female.

Cameroon); Clausen, 1956:227, pl. 1, figs. 1-2 (comparisons, Nigeria);
Sterba, 1959:613, figs. (descr.; European aquarium trade import); Kahs-
bauer, 1962:159 (listed, Nigeria); Daget and IItis, 1965:185 (Gin key; range);
Thys, 1967:83 Gn key, comparisons, possibly conspecific with ansorgii);
Roman, 1970:147, 149, fig. 66 (in key, descr., Rio Muni); Wheeler,
1975:344, text fig. only (aquarium fish; West Africa).

Syngnathus (Parasyngnathus) Ansorgei: Duncker, 1915:85 (n. comb., mis-
spelling, descr.).

Table 1.—Frequency distributions of trunk, tail, and total rings in species of Enneacampus

Trunk rings Tail rings Total rings
Species 12 13 31 32 33 34 35 36 37 44 45 46 47 48 49 50

ansorgii 6 98 1 6 6 Tf i Pac iy? 4 y 6 6
Kaupi 18 yield |2 1

VOLUME 94, NUMBER 2 471

Table 2.—Frequency distributions of dorsal- and pectoral-fin rays and total subdorsal rings
in species of Enneacampus.

Dorsal-fin rays Pectoral-fin rays Subdorsal rings

Species 22 23 24 2 26 27 28 29 12 #13 «#14 «15 16) 17 5.00 5.25 5.50 5.75 6.00 6.25 6.50

ansorgii Ie Abd 2 25a Ge 28 86.4308 Sy ks) AWE ay
Kaupi 115 Ke Bs 3 10mI9 yd LPO NO 2

Syngnathus olssoni Johnels, 1954:348, 399, fig. 18 (orig. descr.; Farida and
Alikiama swamps, Gambia River); Clausen, 1956:227 (comparisons).

Syngnathus olsonni: Daget and IItis, 1965:185 (misspelling, in key, Gambia
and Guinea); Thys, 1967:83 (in key, possibly conspecific with ansorgii).

Diagnosis.—Rings 12-13 + 31-37 = 44-50; dorsal-fin rays 22-29; pec-
toral-fin rays 11-15 (14 or less in 95% of counts); total subdorsal rings 5.0-
6.5; dorsal-fin origin from middle of last trunk ring to origin of 2nd tail ring,
usually at anterior margin of Ist tail ring (70% of specimens examined).
Proportional data based on 48 specimens 54—-118.5 (x = 87.7) mm SL fol-
low: HL in SL 8.4—10.8 (9.4), snout length in HL 2.1—2.8 (2.5), snout depth
in snout length 2.4—4.0 (3.2), length of dorsal-fin base in HL 0.9-1.3 (1.1),
anal ring depth in HL 2.7-4.7 (3.6), trunk depth in HL 2.2-3.3 (2.6), pec-
toral-fin length in HL 4.4—-6.2 (5.2), length of pectoral-fin base in pectoral-
fin length 1.2—1.7 (1.4). Median dorsal snout ridge of subadults and adults
finely denticulate under x30 magnification. See Tables 1-3 for additional
counts.

Coloration.—Snout with or without irregular brown bars or blotches (Fig.
2); opercle with pale stripe along longitudinal ridge, with brownish diagonal
bars below the ridge or mottled throughout; eye usually with short brown

Table 3.—Geographical variation in frequencies of dorsal-fin rays in Enneacampus ansorgii.

Dorsal-fin rays

Locale 22 23 24 25 26 27 28 29
Gambia D 2
Sierra Leone 1 1
Ghana Z 4 2
Nigeria 1 9 38 24 6
Cameroon 1 1 1
Gabon y)
Zaire 1

Angola 1

472 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Enneacampus kaupi. Lateral and dorsal aspects of head and anterior trunk rings.
From 132 mm SL female (CAS-SU 63059).

bars radiating from pupil; head otherwise plain, mottled or blotched with
brown. Body usually with indications of irregular, diffuse, pale bars crossing
dorsum and upper part of sides on every 4th—Sth ring; lower half of side of
trunk usuaily with a rectangular brown blotch on each ring; side of trunk
and tail otherwise variously mottled, blotched or shaded with brown. Dorsal
fin hyaline distally, the proximal third or more of each fin-ray shaded with
brown; proximal third of pectoral fin blotched or shaded with brown, oth-
erwise hyaline; caudal fin irregularly blotched with brown, the margin usu-
ally pale.

Brood pouch and young.—The brood pouch is developed below 12-17
tail rings in 26 adult males (69.5—128.5 mm SL). One (75.5 mm SL) has a
total of 17 eggs in a single 2-row layer through 12 or 14 pouch-rings, another
(94 mm SL) has one layer of 35 eggs through 14 of 16 pouch-rings. Brood-
pouch young (prolarvae and postlarvae sensu Hubbs, 1943) are unusually
large, and one, representative of 32 postlarvae in an 82.5 mm SL male
(GCRL 15485), measured 17.8 mm TL. When best developed, young com-
pletely fill the brood pouch, the pouch-folds fail to meet on the ventral
midline, and the young are partly exposed (Fig. 2).

Comparisons.—The two species of Enneacampus overlap in meristic fea-
tures (Tables 1-2) but E. ansorgii usually has fewer pectoral-fin rays (mod-
ally, 13 against 16 in kaupi). Enneacampus ansorgii has higher values for
ratios of HL in SL (x = 9.4 against 7.1) and snout length in HL (x = 2.5
against 2.0) and these species also differ in other proportional features (see
diagnoses). The median dorsal snout ridge is finely denticulate in subadult
and adult specimens of E. ansorgii (typically entire in kaupi) and the quad-
rate dark blotches on the trunk rings are replaced by arcuate or semicircular

VOLUME 94, NUMBER 2 473

ocellate spots along the inferior trunk ridge in well-marked specimens of E.
Kaupi.

Limited observations suggest that these species also differ in reproductive
capacity and in the size of brood-pouch young. Seven brooding males of E.
ansorgii (75.5-94 mm SL) contained 17 to 36 brood-pouch eggs or postlar-
vae, whereas there were over 800 prolarvae in a male E. kaupi (141 mm
SL). Several prolarvae and postlarvae from different males of E. ansorgii
measured about 11-18 mm TL; prolarvae of E. kaupi are about 4.4 mm TL.

For practical purposes, these species are most readily distinguished by
obvious differences in relative length of the snout (compare Figs. | and 4).

Types.—The holotype of Syngnathus ansorgii (BMNH 1911.6.1.129) has
the following counts and measurements (mm): rings 13 + 36, dorsal-fin rays
29, pectoral-fin rays 14 x 15, subdorsal rings 0 + 6.5, brood-pouch rings
17, anal-fin rays 3, caudal-fin rays 9, SL 110.5, HL 11.4, snout length 4.9,
snout depth 1.5, length of dorsal-fin base 12.1, anal ring depth 4.2, trunk
depth 4.8, pectoral-fin length 2.3, length of pectoral-fin base 1.9. The nares
open through a single pore bilaterally, the lateral trunk ridge ends without
deflection on the left side but the distal extremity is deflected a little ventrad
on the right, and some larvae are retained in the brood pouch. Boulenger’s
(1910) original counts of 15 + 37 rings and subdorsal rings are in error, but
his illustration (Fig. 73) of the holotype shows 13 trunk rings and about 6.5
subdorsal rings.

The syntypes of Syngnathus pulchellus, conspecific with the holotype of
S. ansorgii, consist of a 128.5 mm SL male (BMNH 1874.6.8.20) and a dried
female (BMNH 1888.12.13.41) with an estimated TL of 100 mm. The dorsal
fin is missing in the male and an accurate count of dorsal-fin rays cannot be
obtained from the female. There are 5.75 subdorsal rings in the male, about
5.5 in the female; both have 9 caudal-fin rays; there are 13 + 34 rings in
both, rather than the described count of 13 + 35. Lateral trunk and tail
ridges are discontinuous on both sides of the male and on the left side of
the female; lateral trunk ridges are not deflected distally. The right side of
the female is anomalous in that the lateral trunk and tail ridges are confluent
and this is the ridge configuration described for the species by Boulenger
(1915). In addition to employing this atypical ridge pattern as a principal
character distinguishing S. pulchellus from S. ansorgii and S. kaupi, Bou-
lenger illustrated this configuration (Fig. 74) on the left side of the female
syntype, although it is present only on the right.

Johnels’ (1954) description of S. olssoni was based on six specimens,
including one male with developed brood pouch, 63~—73 mm TL. Four of
these, including the illustrated male “‘type’’ (now 69.5 mm SL), are cata-
loged as NRM 11151; the fate of the others is unknown. These fish, con-
specific with S. ansorgii, have discontinuous lateral trunk and tail ridges,
the lateral trunk ridge is deflected ventrad only on the right side of a 67.5

474 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

inm SL female, and all have 9 caudal-fin rays. The specimen described as
having confluent lateral trunk and tail ridges on the right side is missing.
Johnels reported 14 trunk rings in all of his material but I count 13 in the
four extant fish. My counts of other meristic features fall within the de-
scribed range.

Variation.—Johnels (1954) and Clausen (1956) noted variation in the lat-
eral body ridges in this species and Clausen questioned the reliability of the
lateral ridge configuration as a taxonomic character. Among 199 configu-
rations examined here, the typical pattern of discontinuous lateral trunk and
tail ridges (Fig. 1) occurred in 196 and these ridges were confluent in two
cases. The lateral trunk ridge was straight in 186, whereas it was deflected
distally to end between the lateral midline and the inferior ridge in 11. In
one instance, the superior trunk and tail ridges were confluent rather than
discontinuous. Unilateral or bilateral variation is rather common in some
pipefishes (e.g. Syngnathus acus, S. schlegeli), but in Enneacampus an-
sorgii, aS in most species, ridge configuration varies in less than 10 percent
of those examined and constitutes an important primary character.

The nares are typically 2-pored bilaterally in most pipefishes but the ma-
jority of E. ansorgii examined have a single opening on each side. This
condition appears similar to that reported for Corythoichthys schultzi, an
Indo-Pacific marine species (Dawson, 1977).

Available data (Table 3) suggest a north-south increase in frequencies of
dorsal-fin rays from Gambia to Angola.

Distribution.—Enneacampus ansorgii is known from rivers, streams, and
swamps from the Gambia River drainage to the Cuanza River, Angola.
There are apparently no definite records of its occurrence in estuarine or
marine habitats.

Material examined.—One hundred and thirteen specimens, 40.5—128.5
mm SL, excluding pouch-larvae.

Holotype.—BMNH 1911.6.1.129 (110.5 mm SL, adult male), Angola,
Quanza (= Cuanza) River at Dondo, cast net, 13 July 1910, W. J. Ansorge.

Other material—_GAMBIA: NRM 11151 (4, 61.5-69.5), including holo-
type (male) and 3 paratypes of Syngnathus olssoni). SIERRA LEONE:
UMMZ 187910 (2, 55-94). GHANA: CAS-SU 63060 (3, 56-76), CAS-SU
63061 (1, 63), CAS-SU 63062 (2, 40.5—70), CAS-SU 64635 (1, 88.5), MCZ
48079 (1, 105). NIGERIA: GCRL 15408 (1, 99.5), GCRL 15485 (3, 81.5-88),
GCRL 15486 (2, 93-100), GCRL 15531 (2, 78-99), GCRL 15532 (2, 101-
109.5), ZMUC P.39478-84 (7, 66-87), ZMUC P.39485 (1, 102), ZMUC
P.39486-90 (5, 83-99), ZMUC P.39491-92 (2, 72-—72.5), ZMUC P.39493-94
(2, 68-76.5), ZMUC P.39495-96 (2, 76-95.5), ZMUC P.39497_-99 (3, 58-84),
ZMUC P.39500-04 (5, 70.5-—95.5), ZMUC P.39505—06 (2, 78-107), ZMUC
P.39507-09 (3, 72-85.5), ZMUC P.39510 (1, 88), ZMUC P.39511 (1, 80),
ZMUC P.39512-17 (6, 54-100), ZMUC P.39518 (1, 70), ZMUC P.39519-21

VOLUME 94, NUMBER 2 475

(3, 57.5—99), ZMUC P.39522—28 (7, 49.5—93), ZMUC P.29529-42 (14, 70.5—
92.5), ZMUC P.39543-49 (7, 80-95.5), ZMUC P.39550-51 (2, 75.5—98.5),
ZMUC P.39552-53 (2, 99.5—105), ZMUC P.39554—-55 (2, 95-105.5), ZMUC
P.39556 (1, 94). CAMEROON: BMNH 1874.6.8.20 (128.5, syntype of S.
pulchellus), MNHN 29.91 (1, 84), MCZ 48149 (2, 66-81). GABON: BMNH
1888.12.13.41 (ca. 100, dried syntype of S. pulchellus), GCRL 16280 (1, 49),
MNHN 06-211 (2, ca. 77-95). ZAIRE: IRSNB 19412 (1, 118.5).

Enneacampus kaupi (Bleeker)
Figs. 3-4

Syngnathus spicifer (not of Ruppell): Kaup, 1856:34 (misident. in part,
Guinea sp. only).

Syngnathus Kaupi Bleeker, 1863:24, pl. 4, fig. 2 (orig. descr., Guinea); Du-
méril, 1870:542, 547 (in key, descr. compiled).

Syngnathus kaupi: Gunther, 1870:174 (descr. compiled); Lonnberg, 1895
(Cameroons); Boulenger, 1912:23 (freshwater lagoon at Chiloango, Bas-
Congo); Boulenger, 1915:86, fig. 72 (in key; descr.; Degama and Chiloan-
go, Congo); Metzelaar, 1919:217 (listed); Fowler, 1936:556 (in key, descr.
compiled, Liberia to Congo); Cadenat, 1950:300 (listed; Guinea, Sierra
Leone, Dahomey); Poll, 1953:251, fig. 102 (descr., Liberia to Congo);
Clausen 1956:227 (comparisons; Boulenger’s (1915) ‘‘Degama’’ should be
Degema, Lower Niger); Kahsbauer, 1962:159 (listed, Nigeria); Daget and
Iltis, 1965:185, fig. 114 Gin key; descr.; Ivory Coast; type loc. given in-
correctly as Ghana); Blache et al., 1970:242, fig. 656 (in key); Roman,
1970: 147, fig. 65 (in key; descr.; Rio Muni).

Syngnathus kaupii: Buttikofer, 1890:480 (emendation, Liberia).

Syngnathus Kaupii: Steindachner, 1894:89 (Liberia).

Syngnathus (Parasyngnathus) Kaupi: Duncker, 1915:85 (n. comb.; descr.;
Liberia to Gabon); Monod, 1927:680 (food item of Galeoides decadac-
tylus; Souelaba, Cameroon).

Diagnosis.—Rings 13-14 + 32-34 = 45-47; dorsal-fin rays 26-28; pec-
toral-fin rays 14-17 (15 or more in 91% of counts); total subdorsal rings 5.5—
6.25; dorsal-fin origin from anterior margin to middle of Ist tail ring, usually
at anterior margin (90% of specimens examined). Proportional data based
on 15 specimens 55-141 @ = 91.1) mm SL follow: HL in SL 6.7-7.5 (7.1),
snout length in HL 1.8-2.1 (2.0), snout depth in snout length 3.2—5.4 (4.5),
length of dorsal-fin base in HL 1.2-1.6 (1.4), anal ring depth in HL 3.4—5.3
(4.4), trunk depth in HL 2.8-3.7 (3.2), pectoral-fin length in HL 5.9-8.0
(6.7), length of pectoral-fin base in pectoral-fin length 1.1—-1.5 (1.3). Median
dorsal snout ridge usually entire under x30 magnification. See Tables 1-2
for additional counts.

Coloration.—Roman (1970) described recently collected material as gen-

476 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

erally dusky with a brick-red abdomen; the head plain in males but with
black bars below eye and on lower part of opercle in females; lower part of
side (of trunk) greyish-yellow, with a golden, black-margined, ocellus on
each ring; tail rings black spotted, sprinkled with gold; caudal fin black,
edged with pale. Somewhat faded specimens (CAS-SU 17148) have the dor-
sum and upper parts of sides of body crossed by about 10 narrow and diffuse
pale bars (spaced ca. 4—5 rings apart); each trunk ring with a dark-margined
arcuate or semicircular brown spot extending dorsad from the inferior ridge;
pouch-plates of male with an alternating series of short pale and brownish
bars; dorsal-fin rays and pectoral fins shaded lightly with brown microchro-
matophores; caudal fin brown, with narrow pale margin.

Brood pouch and young.—The brood pouch is developed below 16-18
tail rings in four brooding males (83-141 mm SL) and an immature male (71
mm SL) has evidence of pouch development below 15 rings. None of the
available males have eggs, but pouch-young are present in three. The largest
and best preserved male (141 mm SL) has coiled prolarvae (ca. 4.4 mm TL),
arranged in about four transverse rows and in two layers, through 15 of the
17 pouch-rings; the number of young is conservatively estimated at 850-—
900.

Comparisons.—See under E. ansorgii.

Holotype.—Bleeker (1863) recorded 14 + 34 rings, 26 dorsal-fin rays, 16
pectoral- and 10 caudal-fin rays for the 173 mm TL, female, holotype of E.
kaupi. This specimen (RMNH 3874) now lacks the caudal fin and part of
the tail, the dorsal fin originates at the anterior margin of the Ist tail ring
and there are 6.25 subdorsal rings. Other counts and measurements (mm)
follow: trunk rings 13, remaining tail rings 28, dorsal-fin rays 26, pectoral-
fin rays 16 x 16, head length 21.4, snout length 10.9, snout depth 2.0, length
of dorsal-fin base 16.2, trunk depth 7.1. Bleeker’s atypical count of 10 cau-
dal-fin rays suggests an error in enumeration or a regenerated fin.

Variation.—Atypical body ridge configurations were not noted in material
examined and meristic data show no evidence of geographic variation. Five
specimens examined had 2-pored nares bilaterally, whereas one had a single
pore on the right side and two on the left. Roman (1970) recorded 14 trunk
rings for two of 16 specimens from the Rio Muni region. This count was 13
in the remainder of Roman’s material and in all specimens examined here.

Distribution.—Present materials show that E. kaupi ranges from Guinea
to Zaire where it has been taken with E. ansorgii in the Banana River. |
There is a record from the Loemé River estuary (Congo) and Boulenger
noted the occurrence of this species in brackish water, but most collections
are apparently from freshwater. The only record of E. kaupi from marine
waters is apparently that of Bas (1974) and Lloris and Rucabado (1979). I
have been unable to obtain this specimen, trawled in 24 meters off Punta
Durnford (23°12'N, 16°20’W), and its identity requires verification. |

VOLUME 94, NUMBER 2 Aaa

Material examined.—Nineteen specimens, 55— ca. 165 mm SL, excluding
pouch-larvae.

Holotype.—RMNH 3874 (now 152.5 [orig. ca. 165 mm SL], damaged
female), Guinea.

Other material.—LIBERIA: CAS-SU 48408 (3, 55-77), RMNH 5401 (1,
79.5). GHANA: CAS-SU 63059 (2, 132-141), USNM 219173 (2, 78-99.5).
NIGERIA: BMNH 1902.11.10.301 (1, 68.5). CAMEROON: CAS-SU 17148
(2, 73.5—101), GCRL 17011 (1, 98.5). CONGO: MNHN 1967-185 (1, 110.5),
MNHN 1967-186 (1, 63.5). ZAIRE: AMNH 17135 (1, 83), BMNH
1912.4.1.475-6 (2, 97-119), IRSNB 8701 (1, 84.5).

Acknowledgments

I thank the curators of the various repositories for loans of material and
other courtesies. Special acknowledgment is due M. Boeseman (RMNH)
and A. C. Wheeler (BMNH) for permission to examine type material in
their care. I also thank Bo Fernholm (Roskilde Univ., Roskilde, Denmark)
for obtaining type material of Syngnathus olssoni for my examination. Gift
or exchange specimens were received from W. N. Eschmeyer and T. R.
Roberts (CAS) and J. Nielsen (ZMUC). Drawings are by Mrs. Nancy Gor-
don (GCRL).

Literature Cited

Bas, C. 1974. Distribucion de especies dermersales recogidas durante la expedicion oceano-
grafica “‘Sahara I.’-—Res. Exp. Cient. B/O Cornide 3:187-247.

Blache, J. 1962. Liste des poissons signalés dans |’ Atlantique tropico-oriental sud du Cap des
Palmes a Mossamédeés (province guinéo-équatoriale).—Cah. ORSTOM, ser. Pte. Noire.
2:13-102.

Blache, J., J. Cadenat, and A. Stauch. 1970. Clés de détermination des poissons de mer
senegalés dans |’ Atlantique oriental (entre 20°N et 15°S).—Faune Tropicale, ORSTOM.
18: 1-479.

Bleeker, P. 1863. Mémoire sur les poissons de la cote de Guinée.—Nat. Verh. Holl. Maatsch.
Wetensch. 2(18):1-136.

Boulenger, G. A. 1910. On a large collection of fishes made by Dr. W. J. Ansorge in the

Quanza and Bengo Rivers, Angola.—Ann. Mag. Nat. Hist., 8th ser. 6:537-561.

. 1912. Poissons recueillis dans la region du Bas-Congo par M. le Dr. W. J. Ansorge.—

Ann. Mus. Congo Belge, Zool. ser. 1. 2:1—27.

. 1915. Catalogue of the fresh-water fishes of Africa in the British Museum (Natural

History).—London. 3:1-—526.

Buttikofer, J. 1908. Reisebilder aus Liberia.—E. J. Brill, Leiden. 2:480.

Cadenat, J. 1950. Poissons du mer du Sénégal.—Initiations Afr., IFAN 3:1-345.

Claussen, H. S. 1956. Biological and taxonomical notes on Nigerian fresh-water Syngnathus
(Linné 1758) Kaup 1856, with remarks on the taxonomic value of crista media trunci
and c. superior caudae.—Vidensk. Medd. fra Dansk naturh. Foren. 118:225-234.

Daget, J., and A. Iltis. 1965. Poissons de Cote d’Ivoire (eaux douces et saumatres).—Mem.
IFAN 74: 1-385.

Dawson, C. E. 1977. Review of the pipefish genus Corythoichthys with description of three
new species.—Copeia 1977:295-—338.

478 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

_ 1978. Review of the Indo-Pacific pipefish genus Bhanotia, with description of B. nuda

n. sp.—Proc. Biol. Soc. Wash. 91:392—407.

_ 1979. Review of the polytypic doryrhamphine pipefish Oostethus brachyurus (Bleek-

er).—Bull. Mar. Sci. 29:465—480.

Dawson, C. E., and R. A. Fritzsche. 1975. Odontoid processes in pipefish jaws.—Nature
257:390.

Dumeéril, A. 1870. Histoire naturelle des poissons ou ichthyologie générale. Tome second.
Ganoides, dipnés, lophobranchs.—Paris. 624 pp.

Duncker, G. 1915. Revision der Syngnathidae.—Mitteil. Naturh. Mus. Hamburg 32:9-120.

Fowler, H. W. 1919. The fishes of the United States Eclipse Expedition to West Africa.—

Proc. U.S. Nat. Mus. 56:195-292.

_ 1922. Notes on hemibranchiate and lophobranchiate fishes.—Proc. Acad. Nat. Sci.

Phila. 73:437—448.

_ 1936. The marine fishes of West Africa.—Bull. Amer. Mus. Nat. Hist. 70:1—-605.

Fritzsche, R. A. 1980. Revision of the eastern Pacific Syngnathidae (Pisces: Syngnathiformes),
including both recent and fossil forms.—Proc. Cal. Acad. Sci. 42:181—227.

Giinther, A. 1870. Catalogue of the fishes in the British Museum.—Taylor and Francis, Lon-
don. Vol. 8, 549 pp.

Herald, E. S. 1959. From pipefish to seahorse—a study of phylogenetic relationships.—Proc.
Cal. Acad. Sci. 29:465-473.

Hubbs, C. L. 1943. Terminology of early stages of fishes.—Copeia 1943:260.

Johnels, A. G. 1954. Notes on fishes from the Gambia River.—Arkiv. f. Zool., Stockholm
6:327-411.

Kahsbauer, P. 1962. Beitrag zur kenntnis der Fischfauna von Nigeria.—Ann. Naturhist. Mus.
Wien 65: 139-165.

Kaup, J. 1856. Catalogue of lophobranchiate fish in the collection of the British Museum.—
Taylor and Francis, London. 76 pp.

Lloris, D., and J. Rucabado. 1979. Especies ictiologicas de las expediciones pesqueras real-
izadas en el platforma del NW de Africa (1971-1975).—Res. Exp. Cient. B/O Cornide
8:3—151.

Lonnberg, E. 1895. Notes on fishes collected in the Cameroons by Mr. Y. Sjostedt.—Ofvers.
svenska Vetensk Akad. Handl. 3:179-195.

Metzelaar, J. 1919. Report on the fishes collected by Dr. J. Boeke in the Dutch West Indies
1904-1905, with comparative notes on marine fishes of tropical west Africa.—A. H.
Kruyt, Amsterdam. 315 pp.

Monod, T. 1927. Contribution a la faune du Cameroun. Pisces I.—Faune Col. Fr., Paris.
1:643-742.

Poll, M. 1953. Poissons III. Téléostéens malacoptérygiens.—Res. Sci. Exped. oceanogr. Belg.
eaux cot. afr. Atl. Sud (1948-49) 4(2):1-258.

Roman, B. 1970. Peces de Rio Muni, Guinea Ecuatorial (Aguas dulces y salobres).—Fund.
La Salle Sci. Nat., Barcelona. 295 pp.

Steindachner, F. 1894. Die Fische Liberia’s.—Notes Leyden Mus. 16:1-—96.

Sterba, G. 1959. Freshwater fishes of the world. Revised edn.—Longacre Press, London. 878
pp.

Thys van der Audenaerde, D. F. E. 1965. List of the freshwater fishes presently known from

the Island of Fernando Poo.—Bonn. Zool. Beitr. 16:316—317.

_ 1967. The freshwater fishes of Fernando Poo.—Verh. Kon. v. Acad. Wetens., Belge

29: 1-167.

Wheeler, A. C. 1975. Fishes of the world.—MacMillan Publ. Co., Inc., New York. 366 pp.

Gulf Coast Research Laboratory Museum, Ocean Springs, Mississippi
39564.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 479-482

GONODACTYLUS SIAMENSIS, A NEW STOMATOPOD
CRUSTACEAN FROM THAILAND
Raymond B. Manning and Marjorie L. Reaka

Abstract.—The third shallow water member of the G. falcatus group of
species from Thailand is recognized. It can be distinguished in the field by
its color pattern.

Among the stomatopods collected by one of us (M.L.R.) in Thailand in
1973 was a small species of the Gonodactylus falcatus group which could
be distinguished in the field from the two more common species of the group
found there, G. mutatus Lanchester, 1903, and G. ternatensis De Man,
1902 (see Dingle, Caldwell, and Manning, 1977; Manning, 1978). This
species is described here.

We thank Roy L. Caldwell for help in collecting these specimens; studies
in Thailand were carried out under NSF grant GB-36046. The illustrations
were prepared by Lilly King Manning. All of the specimens have been
deposited in the collections of the Smithsonian Institution under USNM
catalog numbers.

Gonodactylus siamensis, new species
Fig. |

Gonodactylus falcatus.—Reaka, 1979a:238, 252, fig] 197962330), 333), fies
3. [Not Gonodactylus falcatus (Forskal, 1775).]

Material.—Thailand: Gulf of Thailand, Sattahip [12°40’/N, 100°52’E]; in-
tertidal coral rubble reef flat exposed at low tide: 2, 4 July 1973; M. L.
Reaka, and R. L. Caldwell, leg.: 10 3, 21-39.5 mm, 13 2, 16-38 mm CQ.
36 mm long, holotype, USNM 181673; remainder of specimens are para-
types, USNM 181674).

Description.—Rostral plate longer than broad, median spine relatively
long; anterior margin of plate sloping anteriorly or perpendicular to body
line; basal part of plate short, lateral margins divergent, anterolateral angles
acute but broadly rounded. Ocular scales small, breadth of each no greater
than width of rostral spine at base. Anterior 5 abdominal somites lacking
transverse grooves, sixth somite with 6 carinae, variously inflated, usually
unarmed posteriorly in females, occasionally unarmed in small males also:
median carinule absent. Small black spot faintly indicated on each side of
sixth somite between submedian and intermediate carinae. Abdominal
width/carapace length index ranging from 875 in smallest to 780 in largest

480 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Gonodactylus siamensis: a-c, Female, 25 mm long; d-f, female, 36 mm long. a,
d, Rostral plate and ocular scales; b, e, Sixth abdominal somite and telson; c, f, Uropod,

ventral view.

VOLUME 94, NUMBER 2 481

specimens. Telson with length and width subequal or length greater. Dorsal
carinae of telson inflated, especially in males, median and accessory medi-
ans each usually with apical spinule flanked ventrally by rounded excava-
tion; dorsal carinae slender in juveniles, more inflated in adults. Knob dis-
tinctly bilobed. 3 pairs of marginal teeth present, all relatively slender in
females, submedians with movable apices, intermediates and laterals usually
with slender, sharp apices. Intermediate denticles sharp, distinct. Anterior
surface of telson with faint indication of dark spot on each side anterior to
anterior submedian carina. Ventral surface of telson with low postanal ridge
and longer, sharper carina on each submedian tooth. Uropodal endopod
with single line of marginal setae, exopod with 10-13, usually 11, graded
movable spines on outer margin of proximal segment; basal prolongation
with 1 lobe proximally on inner margin of outer spine.

Color in life. —Females uniform olive or chocolate brown, or green, fre-
quently speckled; meral spot yellow with light brown infusion; 4 black spots
on sixth abdominal somite and telson. Males black, dark green, or olive
green, with red posterior edges on abdominal somites. Antennal scales blue;
setae blue proximally, red distally.

Measurements.—Total lengths of males 21 to 39.5 mm, of females 16 to
38 mm. Other measurements, in mm, of female holotype: carapace length
9.0; rostral plate length 2.8, width 2.6; fifth abdominal somite width 7.3;
telson length 6.3, width 6.0.

Remarks.—Two other species of the G. falcatus group of species are
known to occur in Thailand (Dingle, Caldwell, and Manning, 1977; Naiya-
netr, 1980). Gonodactylus ternatensis, the largest of the three species, at-
taining a total length in Thailand of 87 mm, can be distinguished immediately
from G. mutatus and G. siamensis by its long rostral spine and, slender,
sharp carinae as well as an undivided knob on the telson. In life it agrees
with G. siamensis and differs from G. mutatus in having red intersegmental
bands on the body as well as blue antennal scales in the male; it differs from
both species in having an orange rather than a yellow meral spot. Gondac-
tylus siamensis and G. mutatus are of similar size and are superficially
similar morphologically. In life, they can be distinguished immediately by
the blue antennal scale in the former, a red, orange, or yellow antennal scale
in the latter; in addition, G. mutatus lacks the posterior red bands on the
body segments.

Gonodactylus siamensis differs from G. mutatus in the shape of the ros-
tral plate, with the apical spine longer and the basal portion shorter, and the
telson is usually longer, with longer, slenderer marginal teeth in the new
species.

In these specimens, the abdominal width/carapace length index shows a
wide variation between young and old specimens, as follows:

482 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Index
Carapace No. of
length, mm specimens Range Mean
4 1 — 875
5 3 833-860 844
6 yy 833-841 837
7 6 819-877 854
8 1 — 852
9 1 778-826 808
10 2 776-784 780

Many of the specimens of G. siamensis have damaged spines and carinae
on the sixth abdominal somite and telson (Fig. le), suggesting that they are
somewhat aggressive.

Etymology.—The specific epithet is derived from the old name for Thai-
land, Siam.

Literature Cited

Dingle, Hugh, Roy L. Caldwell, and Raymond B. Manning. 1977. Stomatopods of Phuket
Island, Thailand.—Phuket Marine Biological Center, Research Bulletin 20: 1-20, figures
1-11.

Manning, Raymond B. 1978. Notes on some species of the Falcatus Group of Gonodactylus
(Crustacea: Stomatopoda: Gonodactylidae).—Smithsonian Contributions to Zoology
258:1-—15, figures 1-13.

Naiyanetr, Phaibul. 1980. Stomatopoda of Thailand.—95 pages, plates 1-35. Chulalongkorn
University, Bangkok.

Reaka, Marjorie L. 1979a. The evolutionary ecology of life history patterns in stomatopod

Crustacea, pages 235-260. In S. E. Stancyk, editor, Reproductive Ecology of Marine

Invertebrates.—Belle W. Baruch Library of Marine Science, University of South Car-

olina Press.

.1979b. Patterns of molting frequencies in coral-dwelling stomatopod Crustacea.—Bi-

ological Bulletin 156:328-342, figures 1-3.

(RBM) Department of Invertebrate Zoology, National Museum of Natural
History, Smithsonian Institution, Washington, D.C. 20560; (MLR) Depart-
ment of Zoology, University of Maryland, College Park, Maryland 20742.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 483-491

A NEW DEEP-SEA LEECH, BATHYBDELLA SAWYERI,
N. GEN., N. SP., FROM THERMAL VENT AREAS
ON THE GALAPAGOS RIFT

Eugene M. Burreson

Abstract.—Bathybdella sawyeri was common in samples of invertebrates
collected by DSRV Alvin from 2400 meters at 00°48.3’N, 86°13.5’W during
February 1979. The leech is not known to exceed 11 mm in length and has
the following morphological characteristics: tegument smooth, lacking gills
and pulsatile vesicles; eyes and ocelli absent; caudal sucker small; 6 pairs
of testisacs; deep, paired lateral invaginations in XII connected via vector
tissue to large bilobed spermatheca in XII; postceca present.

The discovery of unique biological assemblages near submarine thermal
vents during the first DSRV Alvin expedition to the Galapagos Rift in 1977
(Corliss and Ballard, 1977; Lonsdale, 1977; Corliss et al., 1979) inspired the
Galapagos Rift Biology Expedition in early 1979 (Ballard and Grassle, 1979;
Grassle et al., 1979). Included in the material collected on this expedition
was a small species of piscicolid leech. None of the specimens collected
was from a fish, but the crop of most individuals was filled with red blood
cells indicating that the leech is a fish parasite. Specimens examined include
10 individuals preserved in situ on the outer surface of a single vestimen-
tiferan tube collected on Alvin dive 884 (00°47.7’N, 86°07.7'’ W, 2482 m, 2°C,
25 January 1979), and 46 specimens, ranging from 3 to 11 mm in length,
from the bottom of a small aquarium used to temporarily hold a clump of
mussels collected on Alvin dive 890 (00°48.3'N, 86°13.5’'W, 2447 m, 2°C, 15
February 1979). Five individuals were sectioned (frontal, entire <2; sagittal,
entire; transverse, entire; transverse, anterior half), two wholemounts were
prepared and stained with Semicohn’s acetocarmine, and two specimens
were prepared for examination under the scanning electron microscope.

Bathybdella, new genus

Diagnosis.—Size small, not known to exceed 11 mm; body elongate, cy-
lindrical; paired deep, lateral pits in XII giving appearance of distinct trach-
elosome/urosome division; tegument smooth, lacking papillae, tubercles,
gills, and pulsatile vesicles; caudal sucker slightly subterminal, not wider
than maximum body width; oral sucker well developed; midbody segments
3(9?) annulate; crop strongly compartmented, postceca present; intestine

484 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

FlLOoOmm—H

Figs. 1-6. Bathybdella sawyeri: 1, Reconstruction of digestive system, reproductive sys-
tem, body shape and sucker shape from dorsal aspect; 2, Caudal sucker, lateral view; 3, Male
and female reproductive systems, ventral view; 4, Male and female reproductive systems,
lateral view; 5, Transverse section of terminal portion of male reproductive system at level of |
male gonopore; 6, Spermatheca system, dorsal view. A, anus; AG, accessory gland cells; C,
crop cecum; FP, female gonopore; I, intestine; LI, lateral invagination; M, mouthpore; MP,
male gonopore; O, ovisac; OD, oviduct; P, proboscis; PC, postceca; R, rectum; S, sperma-
theca; T, testisac; VD, vasa deferentia.

VOLUME 94, NUMBER 2 485

Figs. 7-12. Transverse paraffin sections of B. sawyeri illustrating female reproductive sys-
tem. 7, X75; 8-12, 110. 7, Immediately posterior to ganglion in XII showing large lateral
invaginations, LI, and mass of vector tissue, VT; 8, Posterior portion of XII, ventral % of
body, illustrating cords of vector tissue, VT, posterior extensions of lateral invaginations, P,
and common oviduct. CO; 9, Anterior portion of XIII showing bilobed spermatheca, S, with
connecting element, terminal portion of posterior extension of left lateral invagination, P, and
common oviduct, CO; 10, Anterior portion of XIII with paired oviducts, OD, in contact with
ventral portion of spermathecal lobes, S; 11, At ganglion in XIII showing enlarged oviducts,
OD, packed with sperm, and connections (arrows) between oviducts and spermathecal lobes, S;
12, Posterior portion of XIII illustrating oviducts, OD, and testisacs, T.

lacking symmetrical diverticula; 6 pairs of testisacs; bursa moderately large;
deep, paired lateral invaginations immediately posterior to ganglion in XII
opening externally into lateral pits; posterior margins of invaginations cov-
ered by vector tissue with short, paired cords leading to large bilobed sper-

486 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 13. Scanning electron micrograph of B. sawyeri, <120. LI, lateral invagination; MP,
male gonopore; S, sensillae.

matheca (seminal receptable) in XIII; oviducts in contact with ventral sur-
face of spermatheca; marine.

Type-species.—Bathybdella sawyeri n. sp.,

Etymology.—From the Greek, bathy, deep + bdella, a leech.

Bathybdella sawyeri, n. sp.
Figs. 1-13

Diagnosis.—With the characters of the genus; average total length of
mature individuals including suckers, 8 mm (range 6-11 mm); no eyes or
ocelli on either sucker or on body; no pigmentation apparent on body or
suckers; minute sensillae dorsally and ventrally on primary annuli of each
segment on trachelosome and urosome; mouthpore centrally located in oral
sucker; posterior portion of proboscis bulbous; esophageal diverticula ab-
sent; crop ceca trilobed; postceca fused with 4 wide fenestrae; intestine
sinuous with small chambers, rectum large; accessory gland cells covering
dorsal portions of ejaculatory ducts, artrial conua and bursa; bursa confluent
ventrally with specious cavity opening through large male gonopore.

Holotype. —USNM 65773; Paratypes.—USNM 65774, 65775.

Type-locality.—Galapagos Rift, Eastern Pacific Ocean, 00°48.3’N,
86°13.5'W, 2,447 m.

Hosts.—Unknown, but assumed to be deep-sea demersal fishes.

VOLUME 94, NUMBER 2 487

Etymology.—Named in honor of Dr. Roy T. Sawyer in recognition of his
many contributions to hirudinology.

External characters (Figs. 1, 2, 13, measurements of holotype).—Body
elongate, cylindrical, and indistinctly divided into trachelosome and uro-
some, although deep lateral pits at ganglion in XII giving appearance of
distinct separation. Average total length of mature individuals including
suckers 8.0 mm (range 6-11 mm, holotype 9.0 mm). Mouthpore centrally
located in well-developed discoid oral sucker 0.8 mm in diameter eccentri-
cally attached to trachelosome. Oral sucker devoid of eye spots and pig-
ment. Trachelosome tapering slightly toward anterior end where first 2 nu-
chal annuli constricted. Lateral margins otherwise roughly parallel except
for deep lateral pits at level of ganglion in XII where invaginations open
externally. Trachelosome width here narrowing to 0.5 mm (Figs. 1, 13).
Posterior to lateral pits trachelosome widening abruptly into prominent
shoulders, especially when crop is full, and merges with urosome. Male
gonopore a large spherical opening 0.1 mm in diameter (Fig. 13) separated
by 3 annuli from openings to invaginations in lateral pits (Fig. 13). Female
gonopore never observed with certainty even under scanning electron mi-
croscope, but based upon sectioned material should be in same annular
furrow as lateral invaginations or slightly caudad. Urosome widest (1.3 mm)
at midlength of body and tapering to 0.7 mm at caudal sucker, lacking
papillae, tubercles, gills, pulsatile vesicles, pigment and ocelli. Midbody
segments basically 3-annulate, each deeply furrowed primary annulus fur-
ther subdivided such that there appear to be 3 or 4 secondary annuli in
each primary annulus. Minute sensillae (Fig. 13) dorsally and ventrally on
each primary annulus of urosome and trachelosome. Caudal sucker small
(0.8 mm diameter), slightly subterminal, not wider than greatest body width,
and lacking pigmentation and ocelli.

Coelomic system.—Material inadequate for detailed characterization.
Lateral extensions of ventral sinus at ganglia and of dorsal sinus interseg-
mentally in testicular region. Presence of lateral sinuses unconfirmed. Pul-
satile vesicles absent.

Central nervous system.—Central nervous system typical of that of other
piscicolid leeches with ventral nerve cord consisting of anterior ganglionic
mass, 21 segmental ganglia in VII though XXVII, and posterior ganglionic
mass.

Digestive system (Fig. 1).—Mouthpore approximately centrally located
in oral sucker. Proboscis, 530 wm in length, extending to ganglion in IX;
anterior portion about 100 um in diameter, expanding somewhat abruptly
about mid-length into bulbous portion 210 um in diameter (Fig. 1). Bulbous
portion appearing too large to pass through anterior ganglionic mass, unless
quite compressible. Salivary glands located between ganglia in VII and IX.
Crop divided into 2 compartments in trachelosome; first with 2 short anterior

488 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

projections around base of proboscis. No esophageal diverticula present.
Crop lumen expanding between testisacs into large trilobed ceca (Fig. 1).
Intestine and postceca originating immediately posterior to ganglion in XIX.
Intestine a large chamber with short anterior pouches at XIX/XX, narrowing
abruptly at ganglion in XX, becoming tubular, sinuous, with small pouches
in anterior portion of XX, XXI and XXII, the latter displaced laterally to
left. Intestine entering a large, uniformly tubular rectum at XXIII/XXIV.
Posteca fused with 4 wide fenestrae immediately posterior to ganglia from
XX to XXIII. Trilobed nature of crop persisting in postceca to XXIII;
postceca terminate at XXVI.

Reproductive System.—(Figs. 4—13). Six pairs of large testisacs located
intersegmentally in XIII/XIV through XVIII/XIX. Vasa deferentia enlarged
in XIII and entering loosely coiled epididimides in anterior portion of XIII,
continuing cephalad and becoming confluent with ejaculatory bulbs as walls
of ducts become increasingly thicker and more glandular. Immediately pos-
terior to ganglion in X, ducts bending ventrad and lumina becoming very
small. Ejaculatory ducts entering atrial cornua on their anteroventral margin
(Figs. 3, 4). Lumina of atrial cornua merging into small common atrium
which opens into moderately large bursa. Bursa continuing caudad to near
ganglion in XII, bending ventrad and entering a spacious chamber between
body wall and elongated glandular mass of atrium (Figs. 4, 5). This chamber
opening externally through large male gonopore in anterior portion of XII.
Region between ventral surface of ejaculatory bulbs and dorsal surface of
ejaculatory ducts, atrium and bursa covered with large mass of accessory
gland cells (Fig. 3, 4).

Female system having paired, deep, lateral invaginations immediately
posterior to ganglion in XII (Figs. 3, 4, 6). Openings to invaginations ap-
pearing externally as long vertical slits (Fig. 13) in lateral depressions or
pits. These invaginations being spacious, epithelial-lined cavities penetrating
almost to midline of body (Figs. 3, 7). Posterior projections from each in-
vagination terminating near anterior margin of large, bilobed spermatheca
(Fig. 9) located in anterior portion of XIII (Figs. 3, 4, 8, 9). Compact mass
of vector tissue covering dorsal and posterior margins of each lateral inva-
gination and filling area between them (Figs. 6, 7). Vector tissue narrowing
abruptly and bifurcating into 2 short cords of cells prior to merging with
lobes of spermatheca at XII/XIII (Figs. 6, 8). Vector tissue cords without
lumina or surrounding epithelium. Spermatheca, situated in anterior portion |
of XIII, consisting of 2 broadly elongate lobes with a narrow anterior con-
nection. Spermatheca a loose cellular mass bounded by an epithelium except
at extreme anterior portion where a proliferation of vector tissue cord cells
not covered by an external epithelium (Fig. 6). Unlined cavity present in
each spermatheca lobe (Fig. 10). Sperm present in spermatheca of all sec-
tioned leeches, but never present in lateral invaginations.

VOLUME 94, NUMBER 2 489

Ovisacs situated dorsally at XIII/XIV. Left and right oviducts bending
ventrad near first pair of testisacs and continuing anteriorly near body mid-
line (Fig. 12). Oviducts, when contacting ventral surface of spermatheca,
expanding and filled with sperm (Figs. 3, 11). In sectioned material, epithe-
lium of dorsal surface of oviducts and ventral surface of spermatheca be-
coming diffuse such that there are many places where lumina of oviducts
are confluent with spermatheca and sperm can be seen in the process of
entering oviducts (Fig. 11). Oviducts continuing cephalad, still in contact
with spermatheca, then becoming very small tubes (Figs. 3, 10). Oviducts
eventually fusing into common oviduct then bending ventrad and cephalad
to female gonopore somewhat posterior to lateral invaginations (Figs. 4, 8,
9).

Discussion

Bathybdella sawyeri is remarkable for depth of collection, abundance,
and the unique anatomy of the female reproductive system. The depth of
collection is exceeded only by Galatheabdella bruuni from 4400 m and 3880
m in the Tasman Sea (Richardson and Meyer, 1973), and by an unidentified
leech from the deep-sea fish Bassozetus from 3570 m in the Pacific off Costa
Rica listed in the same report. It is likely that the latter leech is Bathybdella
sawyeri. The collection location (9°23'’N, 89°32'W) is near the Galapagos
Rift region and the brief description provided by Richardson and Meyer
(1973), from a single poorly preserved specimen, resembles that of B. saw-
yeri. This suggests that B. sawyeri may not be endemic to vent areas, but
rather may be a parasite of widely distributed deep-sea benthopelagic fishes
such as Bassozetus. The unusually high abundance of the leech in vent
areas may be the result of an abundance of fish hosts attracted by a rich
food supply, and also the presence of hard substrate for cocoon deposition.
According to Cohen (pers. comm.) approximately 16 species of mainly ben-
thopelagic fishes have been photographed, observed, or captured in the
thermal vent areas. Most, including Bassozetus, are rare, but some Ma-
crouridae (benthopelagic) and two species of Zoarcidae (secretive, seden-
tary benthic fishes) are apparently common in the vent area.

Bathybdella sawyeri has one of the most complex arrangements known
in the Piscicolidae for conducting spermatozoa from the external copulatory
zone near the gonopores to the ova. The spermatozoa are apparently intro-
duced into or near the lateral invaginations, migrate through the epithelial
layer of the invaginations to the surrounding vector tissue, and then along
the narrow cords to the spermatheca where they are stored until they pass
into the oviducts. Neither copulation nor attached spermatophores have
been observed in B. sawyeri, however, and it is not known how, or for
certain if, spermatozoa enter the lateral invaginations. An unpaired ventral
invagination, histologically identical to those of B. sawyeri and surrounded

490 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

by vector tissue, occurs in Mysidobdella borealis (Johansson). This struc-
ture often contains spermatozoa and was termed a spermatheca by Burreson
and Allen (1978). It is unlikely that the lateral invaginations in B. sawyeri
function as spermathecae since other structures serve that purpose, and
thus the histologically identical invagination in M. borealis may not function
as a Sspermatheca either. These deep pouches probably hold the spermatozoa
only temporarily. A deep, ventral invagination and a shallow ventral depres-
sion are reported by Brumpt (1900) in Branchellion torpedinis and Cysto-
branchus respirans respectively. While it may be tempting to interpret these
invaginations as relict structures derivable from the paired ectodermal sper-
mathecae of oligochaetes, it is also possible that they are advanced char-
acters derived by invagination of the piscicolid copulatory zone into the
ventral vector tissue mass. The true spermatheca in B. sawyeri is similar
histologically and in segmental position to those of other piscicolids but has
unique features as well. It is strongly bilobed whereas in most piscicolids
spermathecae (used in the broad sense of any large well-defined mass of
vector tissue) are unpaired. These include Marsipobdella sacculata Moore,
Hemibdella soleae Van Beneden & Hesse, Johanssonia arctica (Johans-
son), Calliobdella vivida (Verrill), and Piscicola salmositica Meyer, among
others. However, the basic bilateral nature of the spermatheca in other
species may be reflected by the paired cords of conductive tissue from the
spermatheca to the ovisacs present in H. soleae, C. vivida, P. salmositica,
and others. The primitive leech, Acanthobdella peledina Grube does have
a bilobed spermatheca in XIII and, interestingly, a shallow ventral invagi-
nated pit at XII/XIII (Brumpt, 1900). The most unique feature of the sper-
matheca in B. sawyeri is its position dorsal to the oviducts. In all other
piscicolids the spermatheca or vector tissue mass is situated ventral to the
ovisacs, immediately behind the female gonopore, and closely associated
with the copulatory zone of the ventral body wall. This facilitates transfer
of spermatozoa from spermatophores affixed to the copulatory zone to the
spermatheca. In such arrangements the oviducts usually pass through the
anterior portion of the spermatheca. Even in Marsipobdella sacculata the
elongate, more dorsally situated spermatheca terminates in vector tissue
near the ventral body wall posterior to the female gonopore. In B. sawyeri
the spermatheca is located more in the middle of the body away from the
ventral body wall.

The closest relatives of Bathybdella n. gen. may be Mysidobdella Selen-
sky and Hemibdella van Beneden and Hesse, although neither is really |
very similar. Mysidobdella has a single deep invagination bordered by vec-
tor tissue (Burreson and Allen, 1978), but it is located posterior to the female
pore in XIII while those of B. sawyeri are anterior to the female pore in
XII. Hemibdella possesses a spermatheca and accessory gland cells around
the atrium and lacks eyes and ocelli (Selensky, 1931). Mysidobdella and

VOLUME 94, NUMBER 2 49]

Hemibdella are both small marine leeches with small caudal suckers, but,
unlike B. sawyeri, both have only five pairs of testisacs.

The combination of paired, lateral invaginations; bilobed spermatheca
dorsal to the oviducts; six pairs of testisacs; small caudal sucker, and lack
of pulsatile vesicles clearly separates Bathybdella from all other previously
described genera.

Acknowledgments

I am grateful to Dr. Meredith Jones, USNM, for referring the material to
me, to Dr. Fred Grassle and Linda Morse-Porteous, WHOI, for additional
specimens and collection information, to Dr. Dan Cohen, USNM, for in-
formation on vent area fishes, and to Dr. Roy Sawyer for providing an
unpublished compilation of data on piscicolid spermathecae. This article is
contribution number 22 of the Galapagos Rift Biology Expedition, supported
by the National Science Foundation, and contribution number 993 of the
Virginia Institute of Marine Science.

Literature Cited

Ballard, R. D., and J. F. Grassle. 1979. Return to oases of the deep.—Nat. Geog. Mag.
156:689-705.

Brumpt, E. 1900. Reproduction des hirudinees.—Mém. Soc. Zool. France 13:286—400.

Burreson, E. M., and D. M. Allen. 1978. Morphology and biology of Mysidobdella borealis
(Johansson) comb. n. (Hirudinea: Piscicolidae), from mysids in the western North At-
lantic.—J. Parasitol. 64(6):1082-1091.

Corliss, J. B., and R. D. Ballard. 1977. Oases of life in the cold abyss.—Nat. Geog. Mag.
152:441-453.

Corliss, J. B., J. Dymond, L. I. Gordon, J. M. Edmond, R. P. von Herzen, R. B. Ballard, K.
Green, D. Williams, A. Bainbridge, K. Crane, and T. H. van Andel. 1979. Submarine
thermal springs on the Galapagos Rift.—Science 203:1073-1083.

Grassle, J. F., C. J. Berg, J. J. Childress, J. P. Grassle, R. R. Hessler, H. J. Jannasch, D. M.
Karl, R. A. Lutz, T. J. Mickel, D. C. Rhoads, H. C. Sanders, K. L. Smith, G. N.
Somero, R. D. Turner, J. H. Tuttle, P. J. Walsh, and A. J. Williams. 1979. Galapagos
°79: initial findings of a deep-sea biological quest.—Oceanus 22:2-10.

Lonsdale, P. 1977. Clustering of suspension-feeding macrobenthos near abyssal hydrothermal
vents at oceanic spreading centers.—Deep-Sea Res. 24:857-863.

Richardson, L. R., and M. C. Meyer. 1973. Deep-sea fish leeches (Rhynchobdellae: Pisci-
colidae).—Galathea Rept. 12:113-126.

Selensky, W. D. 1931. Ueber die Gattung Hemibdella nebst einigen allgemeinen Bemerkungen
uber die Organisation der Ichthyobdelliden.—Pubb. Staz. Zool. Napoli 11:1-21.

Virginia Institute of Marine Science and School of Marine Science, Col-
lege of William and Mary, Gloucester Point, Virginia 23062.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 492-502

A NEW DWARF CRAYFISH FROM THE PACIFIC
VERSANT OF MEXICO (DECAPODA: CAMBARIDAE)

Alejandro Villalobos-Figueroa and Horton H. Hobbs, Jr.

Abstract.—Cambarellus prolixus is described from Lago de Chapala, in
the State of Jalisco, Mexico. It shares the lake with two of its closest rel-
atives, Cambarellus chapalanus (Faxon) and C. montezumae (Saussure),
but appears to occupy a different niche. The new species may be distin-
guished from all other members of the genus by the length of the acumen,
which is at least 0.8 as long as, and usually greater in length than, the
remainder of the rostrum.

Introduction

The new crayfish describe? nerein was discovered during the course
of studies on Lago de Chapala, State of Jalisco, conducted by the In-
stituto de Ingenieria of the Universidad Nacional Autonoma de Méx-
ico. Among the organisms collected in the vicinity of Ajijic were several
specimens of a crayfish (acocil) of the genus Cambarellus which were readi-
ly recognized as being different from Cambarellus chapalanus (Faxon,
1898:661), the common crayfish inhabiting much of the littoral zone of the
lake. In subsequent collecting efforts, one of us (AV-F) obtained numerous
specimens of this unique crayfish, verifying its restricted habitat and limited
distribution within the lake.

Cambarellus prolixus, new species
Figs. 1, 2

Diagnosis.—Pigmented, eyes well developed. Rostrum with margins sub-
parallel to concave and bearing spines, acumen at least 0.8 as long as basal
part. Carapace without cervical spine. Areola 3.0 to 4.8 (average 3.7) times
as long as broad and constituting 22.6 to 27.7 (average 26.1) percent of total
length of carapace (39.0 to 48.2, average 44.2, percent of postorbital cara-
pace length). Suborbital angle broadly obtuse to subacute. Branchiostegal
Spine absent. Postorbital ridge with acute cephalic extremity, spine often
Overreaching posterior margin of orbit. Antennal scale approximately 3
times as long as wide, broadest proximal to midlength. Merus of cheliped
with 0-2 dorsal, 1-3 ventral, and 1 distolateral spines. Hooks on ischia of
second and third pereiopods of male, form I, simple, neither overreaching
basioischial articulation nor opposed by tubercle on corresponding basis;
coxa of fourth pereiopod with well developed cephalomesial and caudome-
sial bosses, that of fifth pereiopod subtuberculiform. First pleopods of first

VOLUME 94, NUMBER 2 493

KX

A [a Me ff

hs

: EAS
SAO

k

Fig. 1. Cambarellus prolixus (all illustrations are of holotype except c and h of morphotype,

d, e, and m of allotype, and g of paratypic male, form I): a, Lateral view of carapace; b, c,
Mesial view of first pleopod; d, e, Annulus ventralis; f, Epistome; g, Caudal view of first
pleopods; h, i, Lateral view of first pleopod; 7, Antennal scale; k, Dorsal view of carapace;
1, m, Dorsal view of distal podomeres of right cheliped; n, Lateral view of distal podomeres
of left cheliped; 0, Proximal podomeres of second through fifth pereiopods.

494 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

form male symmetrical, very weakly arched at distal end of proximal third,
lacking both subapical setae and shoulder on cephalic surface; terminal ele-
ments corneous, subparallel in lateral aspect, and directed at about 40 de-
gree angle to main shaft of appendage: mesial process with troughlike groove
mesially, truncate distally, and directed somewhat distolaterally; central
projection tapering from base, its apical part slightly inclined mesially; and
slender caudal process extending caudodistally beyond other 2 elements.
Annulus ventralis about 1.5 times as broad as long, caudal face with con-
spicuous median concavity at base receiving postannular sclerite when an-
nulus rotated posteriorly; prominent undulating horizontal sinus cutting si-
nistral anterior, lateral, and caudal surfaces. Postannular sclerite campanulate
in outline, 1.2 to 1.3 times as broad as long, and base more than 0.75 times
as broad as greatest width of annulus ventralis. First pleopod lacking in
female.

Holotypic male, form I.—Cephalothorax (Fig. la, k) subovate, slightly
compressed. Greatest width of carapace little more than height at caudo-
dorsal margin of cervical groove. Areola about 3.0 times as long as wide
with 5 or 6 punctations across narrowest section, its length constituting 26.1
percent of entire length of carapace (43.4 percent of postorbital carapace
length). Surface of carapace weakly punctate, many punctations bearing
simple setae. Rostrum with slender lateral carinae weakly concave laterally
and terminating in well developed spines overreaching basal segment of
antennule; acumen conspicuously long, overreaching antennal scale, and
extending beyond antennular peduncle by distance subequal to length of
ultimate podomere of latter; dorsal surface concave with usual submarginal
rows of setiferous punctations and scattered ones between. Subrostral
ridges rather weak but evident in dorsal aspect along almost basal third of
rostrum. Left suborbital angle well defined and, although forming obtuse
angle, with small slightly eccentric subacute apex; angle on right side in-
jured. Brachiostegal spine lacking, cephalic extremity of branchiostegite
rounded. Cervical spine absent.

Abdomen slightly narrower than carapace (5.1 and 5.5 mm). Pleura of
third through fifth segments truncate ventrally and lacking angles. Cephalic
section of telson with | spine in each caudolateral corner. Cephalic lobe of
epistome (Fig. 1f) broadly joined to main body, subtriangular, apex of an-
terior angle produced; main body with distinct fovea; epistomal zygoma
rather strongly arched. Proximal podomere of antennule with conspicuous
spine on ventromesial border at about midlength. Antennal peduncle with
well developed spine on distolateral surface of basis; ischium with small
ventral tubercle: flagellum extending caudally to almost midlength of telson.
Antennal scale (Fig. lj) 7.4 times as long as broad, widest proximal to
midlength; mesial margin of lamellar area with 2 subangular bends, proximal
one at widest part of scale and other at about base of distal fourth; disto-

VOLUME 94, NUMBER 2 495

Table 1.—Measurements (mm) of Cambarellus prolixus.

Holotype Allotype Morphotype

Carapace:

Entire length 16.1 14.2 16.9

Postorbital length 9.7 9.0 ¥)53

Width Jeo) S)o5) S)s5)

Height 5:3 5.6 4.9
Areola:

Width 1.4 1.1 1.1

Length 4.2 Sow) 4.3
Rostrum:

Width 1.8 1.8 1

Length 7.0 5.6 8.3

Length of acumen 307) DES Sa
Chela:

Length, palm mesial margin 5.6 3.8 4.0

Palm width 2.8 2.6 2.0

Length, lateral margin 11.3 8.0 9.1

Dactyl length 5.1 4.0 4.7
Abdomen:

Width rll Doll D4

Length 16.8 14.5 16.0

lateral spine overreaching antennular peduncle but falling short of apex of
acumen.

Third maxilliped extending cephalically to slightly beyond basal segment
of antennular peduncle; mesial half of ischium with broad band of stiff sim-
ple setae, single row of plumose setae flanking ventromesial side of lateral
costa, and few between row and mesial band, distolateral angle not pro-
duced; exopod reaching midlength of propodus.

Right chela (Fig. 1/) subovate in cross-section, not strongly depressed;
surface, except for opposable margins of fingers, lacking tubercles and
spines but studded with setiferous punctations, those with short setae much
more numerous than those with long ones; latter more abundant along lateral
surface of fixed finger, on mesial surface and flanking low submedian lon-
gitudinal ridge on dorsal surface of dactyl, and along opposable margins of
both fingers. Opposable margin of both fingers with band of minute denticles
extending along almost entire length; in addition, fixed finger with tubercle
near base, and dactyl with corneous cusp slightly beyond distal end of basal
third of finger (in dorsal aspect, tubercles hidden by minute denticles).

Carpus of cheliped about 1.3 times as long as broad, bearing setiferous
punctations; distal ventrolateral articular area produced in strong acute

496 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

spine. Merus of cheliped (Fig. 11) likewise with setiferous punctations, long-
est setae ventrally and proximolaterally, those situated on latter forming row
of 6 conspicuously long ones; dorsal margin with strong preterminal spine,
ventral surface with 1 at about midlength, and lateral surface with 1 at distal
extremity. Ischium also with setiferous punctations, but without spines or
tubercles; sufflamen clearly defined.

Hooks on ischia of second and third pereiopods (Fig. lo) simple, not
overreaching basioischial articulation and not opposed by tubercle on cor-
responding basis; that on third more tapering and clawlike. Coxa of fourth
pereiopod with conspicuous setiferous caudomesial and cephalomesial boss-
es, latter directly cephalolaterally; coxa of fifth pereiopod with less con-
spicuous tuberculiform caudomesial boss bearing few setae. Sternum be-
tween second, third, and fourth pereiopods rather deep; lateral margins not
strongly produced ventrally but setae borne on them conspicuous.

First pleopods (Fig. 1b, g, i) as described in ‘‘Diagnosis.’’ Lateral lobe
of proximal podomere of uropod broadly rounded, mesial lobe with distinct
spine; distomedian spine on mesial ramus premarginal.

Allotypic female.—Differing, other than in secondary sexual features,
from holotypic male in following respects: rostral margins subparallel to
level of marginal spines, latter more divergent than in holotype, acumen
distinctly shorter, not overreaching antennal scale, dorsal surface with num-
ber of long setae between submarginal rows; in dorsal aspect, subrostral
ridges disappearing beneath lateral carinae on basal fourth of rostrum, sub-
orbital angle lacking subacute apex; abdomen slightly broader than carapace
(5.7 and 5.5 mm); cephalic section of telson with 2 spines in each caudolat-
eral corner; cephalic lobe of epistome triangular, cephalic angle not pro-
duced; mesial borders of antennal scale rounded, 2 subangular bends not so
evident (Fig. 2i), distolateral spines reaching level of apex of acumen; both
fingers of chela (Fig. 1m) with tuft of setae at ventral opposable base, op-
posable margin of fixed finger with tooth at end of proximal third, dactyl
with 2 teeth in corresponding position, minute denticles arranged in single
row on both fingers. See Table | for differences in proportions of chelae and
other body regions.

Annulus ventralis (Fig. le) as described in ‘‘Diagnosis.’’ First pleopods
absent. Basal podomere of uropod as in holotype.

Morphotypic male, form I1.—Differing from holotype in following re-
spects: section of rostrum posterior to marginal spines shorter than in ho-
lotype, acumen overreaching antennular peduncle by almost 3 times length
of ultimate podomere of latter, setae on dorsal surface less numerous than
in holotype; cephalic section of telson with 2 spines in each caudolateral
corner; epistome triangular, cephalic angle not produced; tubercle on is-
chium of antenna acute; flagellum of antenna overreaching caudal margin
of telson; antennal scale with mesial margin of lamella more rounded, sub-

VOLUME 94, NUMBER 2 497

g J

Fig. 2. Cambarellus prolixus: a—c, Dorsal view of cephalic region of paratypic males, form

I; d-g, Lateral view of right merus of paratypic males, form I: h, 7, Dorsal view of right

antennal scale of paratypic males, form I; 7, Same of allotype; k, Diagrammatic representation
of color pattern. Note: Acumen probably regenerated in c.

498 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

angular bends obscured; third maxilliped extending cephalically to base of
ultimate podopere of antennule; opposable margin of fixed finger of chela
lacking tooth, that of dactyl situated on proximal tenth of finger; hooks on
ischia of second and third pereiopods much reduced in size and that on third
tuberculiform rather than clawlike; cephalomesial boss on fourth pereiopod
represented by ridge and not so well defined; first pleopod (Fig. Ic, h) less
inclined caudally, caudal process and central projection disposed as in ho-
lotype but shorter and more robust, mesial process also shorter, and none
of 3 corneous.

Color notes.—The general coloration of this crayfish is grayish brown,
pale to translucent in areas where the chromatophores are widespread or
absent, and dark where they occur in clusters (Fig. 2k). Almost all of the
markings are bilaterally distributed except for the median line on the rostrum
and the broader median cluster on the telson. The dorsomedian line on the
rostrum extends from the apex caudally beyond the base of the orbit. Paired,
laterally convex lines mark the gastric region, and subparallel ones flank the
dorsal part of the cervical groove, that posterior to the groove being darker
than that anterior to it. Concave lines abut the mesial margins of the bran-
chiocardiac grooves, and a pair of conspicuous splotches are present near
the dorsal posterolateral borders of the carapace. Two pairs of short, lon-
gitudinal, darker markings occur on either side of the median line of the
terga of the first and second abdominal segments. The chromatophores on
the third abdominal tergum are, for the most part, rather evenly distributed
but a pair of subcircular spots are present anterolaterally. On the fourth and
fifth terga there are repetitions of the pattern on the first and second, and
the sixth tergum exhibits an anterior dorsolateral pair of curved markings.
The most conspicuous elements of the color pattern on the telson are the
dark clusters at the caudolateral extremities of the cephalic section; the
general distribution of the chromatophores on the remainder of the telson
and uropods is as illustrated. Color on the first pereiopods is concentrated
toward the dorsodistal end of the merus, along the dorsomesial distal part
of the carpus, on and flanking the mesial surface of the propodus, and on
the proximal part of the fingers.

Size.—The largest specimen available is a female having a carapace length
of 17.2 (postorbital carapace length 9.8) mm. Corresponding lengths of the
smallest first form male are 9.9 (6.0) mm, and of the smaller of the two
ovigerous females 11.9 (7.3) mm. |

Type-locality.—Lago de Chapala (Fig. 3), 500 meters from the north levee |
at Ajijic, State of Jalisco, Mexico, at depths of 4 to 5 meters. (See *‘Eco-
logical Notes.’’) |

Disposition of types.—The holotypic male, form I, allotypic female, and —
morphotypic male, form II, are deposited in the National Museum of Nat-
ural History, Smithsonian Institution, nos. 177206, 177207, and 177208, re--

499

VOLUME 94, NUMBER 2

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500 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

spectively. Of the paratypes, 2 dI, 2 dIl, and 2 9 are deposited in each of
the following: Texas Memorial Museum, University of Texas at Austin; Brit-
ish Museum (Natural History); and Rijksmuseum van Natuurlijke Historie.
Of the remaining paratypes, 57d 1, 41d TI, 452, 19 jd, 15 j2, 2 ovig 9,
and |? with young are deposited in the National Museum of Natural History
and 2561, 24dUI, 272, 45 45, 36 j@ in the Instituto de Biologia, Univer-
sidad Nacional Autonoma de México. All except one of the first form males
were collected by the first author and M. E. Zamora on 20 November 1977;
the single male was obtained by C. D. Barbour on 9 December 1979.

Range.—Known only from Lago de Chapala, Jalisco, Mexico.

Variations.—The greatest range of variation noted occurs in the shape of
the rostrum and relative length of the acumen (Fig. 2a—c); the margins may
be subparallel, slightly convergent, or concave, and the acumen ranges in
length from 0.8 to 1.9 times as long as the basal part of the rostrum. Perhaps
most of those rostra in which the acumen is shorter than the basal section
have been injured. The shape of the antennal scale is also variable (Fig.
12h-j), the lateral margin may be almost straight or bowed mesially or lat-
erally; the mesial margin of the lamellate area may exhibit two subangular
bends or be gently rounded in an almost continuous curve. The teeth on the
opposable margins of the fingers of the chelae may vary from 0 to 2 and be
positioned at different levels along the proximal third of the finger. On the
merus of the cheliped (Fig. 2d—g) there may be none to two well developed
premarginal spines dorsally and one to three ventrally. The cephalic section
of the telson bears one or two spines in each caudolateral corner. For other
variations see Table | and ‘‘Diagnosis.”’

Ecological notes.—Lago de Chapala forms a part of the basin of the Rio
Lerma and Rio Santiago which drain some 130,000 km?. It is the largest lake
in Mexico, having an area of 1100 km?, and, in addition to receiving the
waters of the two rivers, it is also fed by runoff from lakeside dwellings,
municipalities, farms, and industries; consequently evidence of eutrophi-
cation and contamination is almost always present.

Prior to the discovery of Cambarellus prolixus, C. chapalanus was the
only crayfish known to occur in the lake (the presence of a third species,
C. montezumae (Saussure, 1857:102), at the eastern end was disclosed dur-
ing the study mentioned in the introductory paragraph above). As shown on
the accompanying map, all three species frequent the marginal or submar-
ginal zone of this body of water; however, C. prolixus does not share the
littoral biotope, but lives at depths of three or four meters where the bottom
is littered with plant debris; some individuals invade depths of seven or
eight meters. The temperature at these depths varies between 18 and 20.5°C;
the pH exhibits values of 8.3 to 8.6; the transparency ranges from 1.2 to 1.4
m; and the oxygen concentration varies from 4.44 to 4.66 ml/I.

Among the animal components of the biocenosis to which C. prolixus

VOLUME 94, NUMBER 2 501

belongs are Anodonta sp. and Pisidium sp. (bivalve mollusks belonging to
the families Unionidae and Sphaeridae, respectively), Planorbis sp. (snails
of the family Planorbidae), larvae and nymphs of various insects including
members of the orders Odonata (Zygoptera and Anisoptera), Hemiptera,
Trichoptera, and Coleoptera (Dytiscidae).

Life history notes.—First form males have been collected on November
20 and December 9. On the former date, two ovigerous females and another
carrying second instar young were obtained. One of the ovigerous females
had a carapace length of 14.6 (postorbital carapace length 9.1) mm and
carried 70 eggs with diameters ranging from 0.8 to 0.9 mm. Corresponding
measurements of the other were 11.9 (7.3) mm and her brood of eggs con-
sisted of 54 with diameters of 0.9 to 1.0 mm. Twenty-one young were found
on a female with lengths of 14.2 (8.4) mm. Inasmuch as the young were in
the second instar, many or a few could have escaped prior to or during
capture and preservation.

Relationships.—Cambarellus prolixus is without doubt more closely al-
lied to the other Mexican members of the genus than to any of those species
occurring in the United States, and shares as much in commion with C.
chapalanus, a species described from the same lake, as with any other
member of the genus. The most striking similarities are in the spination, the
first pleopod of the male, and in the annulus ventralis of the female. More
distantly related are C. montezumae, C. patzcuarensis Villalobos
(1943:607), and C. zempoalensis Villalobos (1943:601). It may be distin-
guished most readily from all other members of the genus in possessing an
acumen that is no less than 0.8 as long as, and usually greater in length
than, the remainder of the rostrum. The spines on the dorsal and ventral
surfaces of the merus of the cheliped (Figs. In, 2d-g) are strikingly more
strongly developed than in any of the four species mentioned except in the
‘giant females’’ (unusually large individuals) of Cambarellus chapalanus.
The ventral spine (never multiple except in C. prolixus and the giant fe-
males) is absent in most C. montezumae, in all C. zempoalensis that we
have examined, and poorly developed in C. patzcuarensis, and the dorsal
one is small to rudimentary in the latter three, and not much better devel-
oped in males and most females of C. chapalanus. The annulus ventralis of
C. prolixus is proportionately broader than in the other four species men-
tioned, but that of C. chapalanus is more similar than are the strikingly
more elongate ones of the remaining three. The mesial process of the first
pleopod of C. prolixus is more robust than that of C. zempoalensis and not
so far removed from the caudal process; also it is shorter than that of C.
montezumae. Whereas C. chapalanus is known to occur in a number of
localities in Jalisco and Michoacan, C. prolixus has been found only in Lago
de Chapala.

Remarks.—Cambarellus prolixus is infested with the same entocytherid

502 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ostracod, Ankylocythere heterodonta (Rioja, 1940), as are all of its relatives
mentioned in the discussion of its relationships (see Hobbs, 1971:33).

Etymology.—Prolixus (L.), stretched out, long; so named because of the
exceedingly long rostrum.

Acknowledgments

We extend our thanks to M. E. Zamora of the Universidad Autonoma
Metropolitana Iztapalapa and to C. D. Barbour of the University of Utah
for their part in obtaining the specimens on which the above description
is based. We are also grateful to Margaret A. Daniel, C. W. Hart, Jr., and
Isabel Pérez Farfante, colleagues at the Smithsonian Institution, for their
criticisms of the manuscript. The assistance of the last in communications
between the authors was invaluable.

Literature Cited

Faxon, Walter. 1898. Observations on the Astacidae in the United States National Museum
and in the Museum of Comparative Zoology, with descriptions of new species.—Pro-
ceedings of the United States National Museum 20 (1136):643-694, plates 62-70.

Hobbs, Horton H., Jr. 1971. The entocytherid ostracods of Mexico and Cuba.—Smithsonian
Contributions to Zoology 81:55 pages, 31 figures.

Rioja, Enrique. 1940. Estudios carcinologicos. V. Morfologia de un ostracodo epizario ob-
servado sobre Cambarus (Cambarellus) montezumae Sauss. de México, Entocythere
heterodonta n. sp. y descripcion de algunos de sus estados larvarios.—Anales del In-
stituto de Biologia, Universidad Nacional Autonoma de México 11 (2):593-609, figures
1-33)

Saussure, Henri de. 1857. Note carcinologique sur la famille des Thalassides et sur celle des
Astacides.—Revue et Magasin de Zoologie Pure et Appliquée, series 2, 9:99-102.

Villalobos Figueroa, Alejandro. 1943. Estudios de los Cambarinos Mexicanos. I. Observa-
ciones sobre Cambarellus montezumae (Saussure) y algunas de sus formas-con descrip-
cidn de una subspecie nueva.—Anales del Instituto de Biologia, Universidad Nacional
Autonoma de México 14 (2):587-611, 2 plates.

(AV-F) Universidad Autonoma Metropolitana Iztapalapa, México 13, D.
F., México, and (HHH) Smithsonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 503-513

CORALANTHURA AND SAURANTHURA, TWO NEW
GENERA OF ANTHURIDEANS FROM
NORTHEASTERN AUSTRALIA
(CRUSTACEA: ISOPODA: ANTHURIDAE)

Gary C. B. Poore and Brian Kensley

Abstract.—Two new anthurid genera, Coralanthura for C. endeavourae
n. sp. and C. ardea n. sp., and Sauranthura for S. goldmanorum n. sp.,
are described from the Great Barrier Reef and the Coral Sea. Coralanthura
is characterized by a 7-articulate maxilliped, non-operculiform pleopods,
and triangular carpi on the posterior pereopods. Sauranthura lacks the sev-
enth pereopod, has a 3-articulate maxilliped, and triangular carpi on the
posterior pereopods.

The anthuridean fauna of temperate southeastern Australia has only re-
cently been the subject of intensive study and has been found to contain
numerous new species and genera (Poore, 1975, 1978). It is not surprising,
therefore, that the fauna of tropical Australia is largely unknown as is the
case for the tropical Pacific (Kensley, 1979). This contribution reports on
two new anthurid genera, one represented by two species. The material
comes from small collections made on reefs in the Coral Sea and on the
Great Barrier Reef. Material is deposited in the National Museum of Vic-
toria, Melbourne (NMV), the Queensland Museum, Brisbane (QM), and the
United States National Museum of Natural History (USNM).

Family Anthuridae
Coralanthura, new genus

Diagnosis.—Eyes present. Antenna | flagellum of 3—5 articles. Antenna
2 flagellum of 6-11 articles. Mandibular palp of 3 articles. Maxilliped of 7
articles, long endite present. Pereopod 1 subchelate, propodus expanded,
larger than pereopods 2 and 3. Pereopods 4—7 with triangular carpi having
short free anterior margins. Pleopod | exopod barely operculiform, not in-
durate. Pleonites 1—5 free, short, subequal in length; 6 fused with telson.
Telson lacking statocysts.

Etymology.—The generic name derives from ‘coral’ for the Coral Sea,
where it was first taken, plus the frequently used name ‘anthura.’

Type-species.—Coralanthura endeavourae, new species.

Remarks.—Only two other genera of anthurids share with Coralanthura
a maxilliped of 7 articles, viz. Neohyssura Amar, and Ocsanthura Kensley.

504 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Coralanthura endeavourae: a, Holotype in dorsal view (scale in mm); b, Antenna
1 5; c, Antenna 2; d, Antenna | 2; e, Mandible; f, Maxilla; g, Maxilliped.

VOLUME 94, NUMBER 2 505

2
f =
=
SS WN

Fig. 2. Coralanthura endeavourae: a, Pereopod 1 2; b, Pereopod 2; c, Pereopod 7; d,
Pereopod 1 6; e, Fused pleonite 6, telson, and left uropod; f, Pleopod 1; g, Pleopod 2 9; h,
Pleopod 2 ¢.

Neohyssura is also similar to Coralanthura in having pleonites 1-5 free,
and pleonite 6 fused with the telson, but differs in being blind, and in having
a poorly developed maxillipedal endite, poor setation on the mandibular
palp, well developed, i.e. expanded pereopods 2 and 3, and a broad prop-

506 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

odus on the posterior pereopods. The blind genus Ocsanthura has relatively
more elongate pleonites, a quadrate carpus of the posterior pereopods, a
less strongly developed maxillipedal endite, a slender acute mandibular mo-
lar and broad lamina dentata, and pleonite 6 free.

Coralanthura endeavourae, new species
Figs. 1, 2

Description.—&. Integument not indurate; with dorsal pigmentation
patches on head, pereon, and pleon. Body proportions: C < 1=2=3 <
4>5>6> 7. Head with blunt rostrum as long as rounded anterolateral
lobes; eyes well pigmented. Pereonites 4-6 each with shallow, elongate
middorsal pit. Pleonites 1-5 free, short, subequal, pleonite 6 fused to telson,
with strong middorsal excavation in posterior margin. Telson proximally
narrow, widest at midlength, tapering evenly to rounded apex, margins un-
evenly serrate, with short lateral setae, and elongate distal setae.

Antenna | flagellum of 5 articles, equal in length to second and third
articles of peduncle together. Antenna 2 peduncle with articles 4 and 5
relatively elongate; flagellum of 11 articles, shorter than 2 proximal peduncle
articles. Mandibular palp of 3 articles, second as long as first and third together;
article 3 with comb of 7 fringed spines, penultimate spine longest; incisor
with single cusp; lamina dentata with 5 serrations; molar bluntly rounded.
Maxilliped of 7 articles, third article short; seventh minute, subterminal;
narrow distally rounded endite reaching to middle of maxillipedal article 5.
Pereopod | subchelate, stout; merus with blade-like spine on anterior mar-
gin; propodus inflated, as deep as long, with straight setose cutting edge;
dactylus strongly hooked, shorter than palm, unguis almost half length of
dactylus. Pereopods 2 and 3 similar, less robust than first, propodi not in-
flated, with blade-like anterior spine; propodus stoutly cylindrical, with se-
tae and distal spine on palm; unguis one-third length of curved dactylus.
Pereopods 4-7 similar, merus with blade-shaped anterior spine; carpus tri-
angular, but with short free anterior margin; propodus with 4 evenly-spaced
spines on posterior margin; dactylus with short unguis. Pleopods subequal
in length and form. Pleopod 1 non-operculiform, endopod as long as exopod,
but less than half width, both rami with distal plumose setae. Uropodal
peduncle short, half as long as telson; exopod lanceolate-acute, about one-
third as wide as long; endopod reaching to end of telson, subovate, both
rami with setose margins.

6. Head with large swollen eyes; antenna | flagellum of 6 articles bearing
dense whorls of aesthetascs. Pereopod | palm setose; pereopod 2 as in
female; pereopod 3 with distally produced carpus. Pleopod 2 with club-
shaped stylet little longer than rami.

Etymology.—The species is named for H.M.S. Endeavour, the ship in
which Captain James Cook visited Northern Australia in 1770.

VOLUME 94, NUMBER 2 507

Material.—Holotype: NMV J821 and J822 (slide), ovigerous 2°, TL 7.4
mm: Coral Sea, cay north of Long Island, Chesterfield Reefs, 19°48’S,
158°17’E; 10 m from reef edge; coll. N. L. Bruce, 10 May 1979.

Paratypes: NMV J823, | non-ovigerous 2 6.0 mm, | manca 3.0 mm, |
6 4.4 mm; QM W8096, 1 ¢ 4.5 mm; QM W8106, 1 non-ovigerous ¢ 7.3
mm, | manca 3.0 mm; from type locality. NMV J824 and J825 (slide), 1 ¢
4.6 mm; USNM 181712, 3 non-ovigerous, 2, 4.2 mm, 4.9 mm, 5.8 mm;
Long Island, Chesterfield Reefs, seaward edge of reef, 15 m; coll. N. L.
Bruce, 6 May 1979. NMV J826, head and pleon only; Queensland, Lizard
Island, 14°40’S, 145°28’E, Great Barrier Reef, outer barrier reef near plat-
form, | m, coral rubble with red encrusting algae; coll. B. Kensley, 29 May
1980.

Coralanthura ardea, new species
Figs. 3, 4

Description.—® . Integument not indurate, with small dorsal spots of pig-
ment on head, posterior margin of pereonites 2-6, and pleonite 6. Body
proportions: C< 1>2>3<4=5<6*>7. Head with broad rostrum as
long as rounded anterolateral lobes; eyes small, well pigmented. Pereonites
4-6 with very shallow dorsal pits. Pleonites 1-5 free, pleonite 6 fused with
telson, with strong middorsal excavation in posterior margin. Telson narrow
proximally, widest at midlength, tapering to rounded apex; distal margins
unevenly serrate; short setae laterally, elongate setae distally.

Antenna 1 peduncle with second and third articles subequal in length,
flagellum of 3 articles, equal in length to last article of peduncle. Antenna
2 peduncle with fifth article twice length of fourth, flagellum of 6 articles,
shorter than last peduncle article. Mandibular palp of 3 articles, second as
long as first and third together; article 3 with comb of 7 spines, distalmost
longest; incisor with 3 cusps; lamina dentata with 5 serrations; molar ob-
solete. Maxilliped of 7 articles, third short, seventh minute, terminal; narrow
apically acute endite reaching almost to end of maxillipedal article 5. Per-
eopod | subchelate, stout; ischium and basis with deep flanges on anterior
margins; propodus almost as deep as long, inflated, palm with 2 proximal
teeth plus setae; dactylus curved, as long as palm, unguis one-quarter length
of dactylus. Pereopod 2 less well developed than first; ischium and basis
with flanges on anterior margin; propodus with large proximal and smaller
distal tooth on palm; unguis one-third length of curved dactylus. Pereopod
3 similar to 2, but toothed palm of propodus shorter. Pereopods 4—7 similar;
carpi triangular, with free anterior margin considerably shorter than poste-
rior margin; propodus with row of about 9 spines along posterior margin;
dactylus with short unguis. Pleopod | approaching operculiform condition,
not indurate; exopod almost twice length of pleopod 2, distally setose; en-
dopod small, triangular, less than half length of exopod, with 2 terminal

508 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Coralanthura ardea: a, Antenna 1; b, Antenna 2; c, Maxilliped; d, Mandible; e,
Pleopod 1; f, Pleopod 2.

setae. Pleopod 2 with subequal rami, distally setose. Uropodal peduncle
short, half length of telson; exopod broadly lanceolate, about twice as long
as greatest width; endopod reaching to end of telson, widest basally, taper-
ing to rounded apex; both rami with setose margins.

Etymology.—The species, taken from Heron Island, is named for Ardea,
a genus of heron.

Material.—Holotype: NMV J827, ovigerous 2, TL 4.8 mm; Queensland,
Heron Island, Great Barrier Reef 23°27'S, 151°55’E; reef flat, coll. N. L.
Bruce, 15 Jan. 1979. Paratypes: NMV J828, 2 non-ovigerous 2, 3.2 mm, 3.9
mm; NMV J829, J830 (slide), 1 2 4.3 mm, QM W8671, 3 non-ovigerous
2, 2.0 mm, 3.5 mm, 3.6 mm; USNM 181713, 2 non-ovigerous ¢ , 3.5 mm,
4.3 mm; all from type-locality.

Remarks.—C. endeavourae and C. ardea are placed in the same genus
as they show a similar structure in the pleonal condition, maxilliped, man-
dible, and pereopods. The structure of pleopod 1 (along with the number of

VOLUME 94, NUMBER 2 509

Fig. 4. Coralanthura ardea; a, Pereopod 1; b, Pereopod 7; c, Pereopod 2; d, Telson; e,
Uropod.

510 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

articles in the first and second antennae, propodus of pereopod 1, and
maxillipedal palp) serve to separate the species, with C. ardea showing
a more advanced condition than C. endeavourae. In the latter, the endo-
pod of pleopod 1 is as long as, but only about half the width of the
exopod, which is broader than in the following subequal pleopods. In
C. ardea, however, pleopod | is longer than the following pleopods, the
endopod is reduced to a short triangular ramus, while the broad exopod
is becoming operculiform.

C. ardea is known only from the type-locality at 23.5°S, considerably
further south than C. endeavourae.

Sauranthura, new genus

Diagnosis.—Eyes present. Antenna | flagellum of 1 or 2 articles. Antenna
2 flagellum of 2 articles. Mandibular palp of 3 articles. Maxilliped of 3 ar-
ticles, endite present. Pereopod 1 subchelate, propodus expanded, larger
than pereopods 2 and 3. Pereopods 4—6 with carpus triangular. Pereopod 7
lacking. Pleonites 1—5 fused, 6 free. Pleopod 1 exopod operculiform. Telson
with 2 basal statocysts.

Etymology.—The generic name is derived from the Greek ‘‘sauros’’—a
lizard, the type locality being Lizard Island.

Type-species.—-Sauranthura goldmanorum, new species.

Remarks.—Among the Anthuridae two genera lack pereopods on the sev-
enth pereonite, viz. Hyssura Norman and Stebbing, 1886, and Exallanthura
Kensley, 1980. The former possesses a 5- or 6-segmented maxilliped, ple-
onites 1—5 free, pleonite 6 fused with the telson, and the exopod of pleopod
1 non-operculiform. The latter genus, with pleonites 1-5 fused, an opercu-
liform exopod of pleopod 1, and a 3-segmented maxilliped, resembles Saur-
anthura more closely, but the single-segmented mandibular palp immedi-
ately separates the Indian Ocean genus from the present material. In general
body form, maxillipedal, antennal, uropodal, and telsonic structure, Saur-
anthura is very similar to Pendanthura Menzies and Glynn, 1967, but the
single-segmented mandibular palp of this latter genus again distinguishes it
from Sauranthura.

Sauranthura goldmanorum, new species
Fig. 5

Description.—Integument thin, non-indurate. Body proportions: C >
1>2=3=4=5>6>7; pereonite 7 about half length of pereonite 6.
Head with rounded rostrum extending beyond rounded anterolateral lobes;
band of red-brown pigment between well-pigmented dorsolateral eyes. Ple-
onites 1-5 fused, with lateral slits indicating segmentation; pleonite 6 free,

VOLUME 94, NUMBER 2 511

Fig. 5. Sauranthura goldmanorum: a, Paratype in dorsal view (scale in mm); b, Antenna
1; c, Antenna 2; d, Mandible; e, Maxilla; f, Maxilliped; g, Pleopod 1; h, Uropodal basis and
endopod; i, Pereopod 1; j, Pereopod 2; k, Pereopod 6.

512 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

with middorsal slit in posterior margin. Telson elongate/ovate, distally
broadly rounded, with 6 terminal setae, broad hyaline margin; 2 statocysts
in proximal half.

Antennal peduncle with first article subequal in length to second and third
articles together, 2 articles of flagellum very short, setose. Antenna 2 pe-
duncle with 3 distal articles subequal in length; flagellum of 2 very short
setose articles. Mandibular palp of 3 articles; middle article longer than first
or third, last with 2 terminal setae; molar rounded, incisor of 3 blunt cusps;
lamina dentata with 6 serrations. Maxilliped of 3 articles, 2 distal articles
subequal in length; thin-walled, narrowly-tapering endite on inner surface.
Pereopod 1 carpus distally rounded; propodus expanded, palm with thin
hyaline margin bearing few short setae. Pereopods 2-6 essentially similar;
carpus with anterior margin shorter than posterior; propodus becoming pro-
gressively more elongate posteriorly, bearing strong serrate spine postero-
distally. Pleopod 1 exopod operculiform, slightly longer than and 3 times
width of endopod; latter with 3 distal plumose setae. Uropodal exopod
ovate, with scalloped hyaline margin bearing plumose setae and few simple
setae, just reaching base of endopod; latter ovate, bearing several elongate
simple setae.

Etymology.—The species is named for Barry and Lois Goldman of Lizard
Island, Great Barrier Reef, Queensland.

Material.—Holotype: NMV J831, 1 non-ovigerous ¢ 2.0 mm; between
Palfrey and South islands, off Lizard Island, 14°40’S, 145°28’E, Great Bar-
rier Reef, from algai turf growing on large dead coral fragments, 3 m, coll.
B. Kensley, 30 May 1980. Paratypes: NMV J832, J833 and J834 (slide), 4
non-ovigerous ?, 1.9-2.2 mm, crest between Palfrey and South Island, off
Lizard Island, 14°40’S, 145°28’E, Great Barrier Reef, coral rubble, 0.5 m,
coll. B. Kensley, 31 May 1980. USNM 181714, 1 non-ovigerous ? 1.8 mm,
lagoon between Palfrey and South islands, off Lizard Island Great Barrier
Reef; Acropora coral rubble, 8-10 m, coll. B. Kensley, 28 May 1980.

Acknowledgments

Our sincere thanks are due to Niel Bruce, University of Queensland, for
collecting some of the material mentioned in this paper; to Barry and Lois
Goldman of Lizard Island, for their assistance and hospitality to the second
author during a short visit in 1980; and to Dr. Thomas E. Bowman of the
Smithsonian Institution for his helpful criticism of the paper.

Literature Cited

Kensley, B. 1979. New species of anthurideans from the Cook and Fiji islands. (Crustacea:
Isopoda: Anthuridea).—Proceedings of the Biological Society of Washington 92:814—
836.

VOLUME 94, NUMBER 2 513

. 1980. Anthuridean Isopod Crustaceans from the International Indian Ocean Expedi-
tion, 1960-1965, in the Smithsonian Collections.—Smithsonian Contributions to Zoology
304: 1-37.

Menzies, R. J., 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.
Norman, A. M., and T. R. R. Stebbing. 1886. On the Crustacea Isopoda of the ‘Lightning’,

‘Porcupine,’ and ‘Valorous,’ Expeditions.—Transactions of the Zoological Society,
London 12:77-141.
Poore, G. C. B. 1975. Australian species of Haliophasma (Crustacea: Isopoda: Anthuridae).—
Records of the Australian Museum 29:503-533.
1978. Leptanthura and new related genera (Crustacea, Isopoda, Anthuridea) from
eastern Australia.—Memoirs of the National Museum of Victoria 39: 135-169.

(GCBP) Department of Crustacea, National Museum of Victoria, Mel-
bourne, Victoria, Australia; (BK) Department of Invertebrate Zoology,
Smithsonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 514-524

CERATOCOLAX MYKTERNASTES, NEW SPECIES
(COPEPODA, BOMOLOCHIDAE) PARASITIC IN
THE NASAL SINUS OF HAEMULON SCIURUS

(PISCES, POMADASYIDAE) FROM BELIZE

Hillary Boyle Cressey

Abstract.—Ceratocolax mykternastes, n. sp., is described from the nasal
sinus of Haemulon sciurus from off Belize. Descriptions of both the female
and male are given. The males differ in one respect from the generic diag-
nosis of Ceratocolax. Placement in the genus Ceratocolax is based primarily
on female structures.

During fieldwork in Belize in March 1980, I discovered a new species of
Ceratocolax Vervoort in the nasal sinuses of the bluestriped grunt, Hae-
mulon sciurus (Shaw). The only other species of Ceratocolax known thus
far is C. euthynni Vervoort, parasitic in the nasal sinuses of the scombrid
fishes Euthynnus alleteratus and Sarda sarda (Vervoort, 1965; Cressey and
Cressey, 1980).

The illustrations were made with the aid of a Wild Drawing Tube. All
measurements are of the holotype or allotype. All material is in the collec-
tions of the Smithsonian Institution.

Ceratocolax mykternastes, new species

Material examined.—Holotype 2 (USNM 181890), allotype 6 (USNM
181891), and 11 2,3 3 paratypes (USNM 181892) collected from the nasal
sinuses of 24 Haemulon sciurus from off Carrie Bow Cay, Belize, 4-9 March
1980. In addition, one female and one male were dissected and illustrated
and 2 females were prepared for examination with a scanning electron mi-
croscope; these additional specimens were from the same collection as the
type-material.

Female.—Body form as in Figure 1. Total length 1.94 mm, greatest width
1.16 mm (measured at widest part of cephalon), length of cephalon 0.58

—>

Figs. 1-8. Ceratocolax mykternastes, female: 1, Dorsal; 2, Genital segment, ventral; 3,
Genital segment and abdomen, dorsal; 4, Last abdominal segment and caudal rami, ventral;
5, First antenna; 6, First antenna spine; 7, Cephalon and first antenna, lateral; 8, Mandible,
paragnath, first maxilla, second maxilla.

515

VOLUME 94, NUMBER 2

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516 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

mm; cephalon with median, longitudinal groove. Genital segment (Fig. 2)
wider than long (265 x 324 um), ventrally with prominent patches of spi-
nules as indicated in figure; spinules may appear as one patch, or there may
be more or less distinct areas between patches; in all specimens examined,
however, outline of spinulose area was similar. Abdomen (Fig. 3) 3-seg-
mented, segments measure (I X w) 123 x 188 um, 118 x 165 um, and
106 x 153 um respectively; last segment (Fig. 4) ventrally with 2 prominent
patches of large spinules. Caudal rami (Fig. 4) slightly longer than wide
(70 x 53 wm); each ramus with 6 setae, 2 much longer than other 4, and
ventral patch of spinules similar to those on last abdominal segment; longest
seta measures 613 wm.

First antenna (Fig. 5) 5-segmented (second segment incompletely divid-
ed), first and second segments together bearing row of 15 stout, plumose
setae, last 2 segments each with one aesthete. Arising dorsally near juncture
of first and second segments is a long, curved, sclerotized spine bearing a
small, naked seta; seta is embedded in membrane reaching from tip of spine
to midlength of edge; tip of spine curves mediad toward crenulate cephalic
knob (Figs. 6, 7, 27). Rostrum with 2 ventral hooks (Fig. 28). Second an-
tenna similar to that of C. euthynni, with numerous rows of small, closely
spaced hooklets, 4 hooked spines, and 2 apical setae. Mandible, paragnath,
first maxilla and second maxilla as in Figure 8; labrum with 2 large patches
of stout spinules. Maxilliped (Fig. 9) with one naked and 3 plumose setae;
maxilliped hook with outer accessory tooth.

Legs 1-4 biramous, rami 3-segmented except leg 1 exopod. Legs 2-4 all
segments, except leg 2 endopod third segment, with patches or rows of stout
spinules along outer or distal edges, as indicated in figures. Leg | (Fig. 10)
coxopod distolateral corner heavily haired, distal edge with short row of
spinules; basipod with long, stout, plumose seta on distolateral corner, ven-
tral surface with large patch of spinules near insertion of endopod, and
smaller rows and patches of spinules as indicated in figure; exopod 2-seg-
mented, second segment incompletely divided, first segment with outer
crenulate spine, second segment with 4 outer spines and 6 terminal to inner
setae; endopod 3-segmented, first and second segments each with one inner

—
Figs. 9-14. Ceratocolax mykternastes, female: 9, Maxilliped; 10, Leg 1; 11, Leg 2; 12, Leg
38 13), Ieee Cle IAN I Lyerer Sy

Figs. 15-22. Ceratocolax mykternastes, male: 15, Dorsal; 16, Genital segment and abdo-
men, ventral; 17, Last abdominal segment and caudal rami, ventral; 18, First antenna; 19,
Second antenna; 20, Mandible, paragnath, first maxilla, second maxilla; 21, Maxilliped; 22,
Leg 1.

Figs. 23-26. Ceratocolax mykternastes, male: 23, Leg 2; 24, Leg 3; 25, Leg 4; 26, Leg 5.

517

VOLUME 94, NUMBER 2

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518 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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VOLUME 94, NUMBER 2

520 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

VOLUME 94, NUMBER 2 521

seta and row of short hairs along distolateral edge; third segment with one
small spine on outer corner, and 5 setae (all setae plumose); interpodal plate
(Fig. 29) with flattened, overlapping spinules. Leg 2 (Fig. 11) coxopod with
short row of spinules on distolateral corner; basipod with slender, plumose
seta on distolateral corner; exopod first segment with outer spine, outer and
inner edge of segment haired, second segment with outer spine and inner
seta, third segment with 3 outer spines and 6 setae, 4 outermost setae with
long plumes on inner edge, short hairs on outer edge (see Fig. 30), all spines
with short hairs along margins and flagellum at tip (see Fig. 31); endopod
somewhat inflated, first segment with stout, inner seta, second segment with
2 inner setae, third segment with 3 inner to terminal setae and 2 outer,
sclerotized spines. Leg 3 (Fig. 12) similar to leg 2 with following exceptions:
exopod first segment outer edge spinulose, third segment with 2 outer spines
and 6 setae; endopod not inflated, last segment with 2 setae and 2 spines,
spines larger than those on leg 2. Leg 4 (Fig. 13) exopod similar to leg 3
except more slender and patches of spinules larger; endopod elongate (es-
pecially third segment), extending well beyond exopod; first and second
segments each with short, inner seta, proximal half of each seta plumose,
distal half spinulose, third segment with distal seta flanked by 2 spinulose
spines. Leg 5 (Fig. 14) uniramous, first segment with outer plumose seta
and outer patch of long, slender spinules; free segment with 3 long, slender,
spinulose spines and one naked seta, segment with several patches of long,
slender spinules. Leg 6 represented by 3 naked setae on genital segment
near area of egg sac attachment (Figs. 2, 3).

Male.—Body form as in Figure 15. Total length 1.25 mm, greatest width
0.50 mm, length of cephalon 0.32 mm. Genital segment (Fig. 16) slightly
longer than wide (230 x 206 wm). Abdomen 2-segmented, segments mea-
sure (1 X w) 88 x 94 um, and 64 x 88 um respectively; last segment (Fig.
17) ventrally with several patches of stout spinules. Caudal rami (Fig. 17)
longer than wide (47 x 29 um), each ramus with 6 setae (2 much longer
than other 4), and ventral patch of stout spinules; longest seta 460 um.

First antenna (Fig. 18) 5-segmented with anterior row of 15 stout, plumose
setae on first and second segments; last 2 segments each with one aesthete;
no spines or modified setae. Rostral hooks absent. Second antenna (Fig. 19)
similar to that of female, but rows of spinules not so closely spaced. Man-
dible, paragnath, first and second maxillae as in Figure 20. Mouthparts very
small, and accessory mandibular blade, present in female, could not be seen
in male specimen dissected and illustrated; first maxilla with minute spinules

<—

Figs. 27-32. Ceratocolax mykternastes, female: 27, First antenna spines, lateral, x 750; 28,
Ventral rostral hooks, x1050; 29, Leg 1 interpodal plate, x775; 30, Leg 2 exopod seta, x1600;
31, Leg 2 exopod spine, <1500; 32, Leg 3 exopod, tip of last segment, «1000.

Sy PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

or hairs on basal segment; paragnath with short row of slender spinules
along distal edge. Maxilliped (Fig. 21) basal segment with one plumose seta;
second segment with patch of short spinules, 2 plumose setae, and row of
stout, triangular spinules; last segment clawlike, with plumose seta and
smaller, naked setule, inner edge with row of small, toothlike spinules.

Legs 1—4 biramous, rami 2-segmented; all segments with patches or rows
of stout spinules along outer or distal edges as indicated in figures. Leg 1
(Fig. 22) coxopod with row of stout spinules along distolateral edge, and
plumose seta on distomedial corner; basipod with long, outer plumose seta,
a somewhat horseshoe-shaped patch of spinules on inner surface and short,
spinulose seta medial to insertion of endopod; exopod first segment with
outer spine; second segment with 4 outer spines, and 4 inner to terminal
plumose setae; proximal 4 spines haired on margins, with flagellum at tip;
distal spine elongate, tapered and sickle-shaped at tip, inner edge plumose,
outer edge with short, closely spaced hairs; endopod first segment with
inner seta, second segment somewhat elongate, with short, outer spine and
6 inner to terminal plumose setae, outer margins of endopod segments heavi-
ly haired; interpodal plate with long, stout spinules. Leg 2 (Fig. 23) coxopod
with row of spinules on outer distal corner; basipod with outer seta smaller
than that on leg 1, small patch of spinules on inner surface; exopod simular
to leg 1 except second segment with 2 rows of spinules on outer edge, and
5 plumose setae; also, distal spine more elongate than that of leg 1; endopod
first segment with inner seta, second segment with 2 outer to terminal spines
and 4 inner to terminal plumose setae. Leg 3 (Fig. 24) similar to leg 2 with
following exceptions: exopod second segment with only 2 short, flagellated,
outer spines; endopod second segment with only 3 plumose setae. Leg 4
(Fig. 25) exopod similar to leg 3 except proximal 3 outer spines slightly more
elongate and serrate rather than haired on margins, second segment with
only 4 plumose setae; endopod first segment with inner seta, second segment
elongate with terminal spinulose seta flanked by 2 elongate spinulose spines.
Leg 5 (Fig. 26) uniramous, basal segment with sparsely plumose seta, free
segment with stout spinules along outer and distal margins, distally with
stout inner spine and outer seta.

Etymology.—The specific name is from the Greek, mykter (nostril), and
nastes (dweller), referring to the habitat of this parasite.

Remarks.—The females of C. mykternastes agree with Vervoort’s (1965)
generic diagnosis of Ceratocolax, especially in the unusual structure of the
first antenna. Ceratocolax mykternastes can be distinguished from C. eu-
thynni by a number of features: C. mykternastes has a prominent spinulose
area on the ventral surface of the genital segment, this is lacking in C.
euthynni, the adults of which have prominent lateral processes on the genital
segment; the dorsal antennal spine of C. mykternastes has an accessory
seta near the tip, this seta is not present on C. euthynni; leg 2 endopod first

VOLUME 94, NUMBER 2 523

and second segments of C. mykternastes have stout spines along the distal
edges, rather than tufts of hairs as in C. euthynni; C. mykternastes leg 4
exopod last segment has 2 spines, 6 setae, while C. euthynni has 3 spines,
4 setae. In addition, there are several other, minor points of difference
between the two species.

Male C. mykternastes differ from Vervoort’s generic diagnosis of Cera-
tocolax in one respect; that is, the number of segments of legs 1-4. The
rami of legs 1-4 of C. euthynni are 3-segmented (except leg 4 endopod
2-segmented); those of C. mykternastes are all 2-segmented. A comparison
of the spine and seta formulae of these two species shows that both species
have the same total number of spines on leg 1 endopod and on both rami
of legs 2-4. The total number of setae has been reduced by 1 in C. mykter-
nastes on both rami of legs 2 and 3, and leg 4 exopod; the total number of
spines and setae is the same for both species for legs | and 4 endopods. In
all other respects, males of the present species agree with the generic di-
agnosis of Ceratocolax.

In spite of this apparently major difference between the males, I have
placed this species in Ceratocolax principally on the basis of the female
morphology. Many copepod species (and genera) are known only from fe-
males, therefore, it would not seem wise, at this time, to erect a new genus
on the basis of a male character alone. Collections of more members of this
genus will perhaps lead to modifications of the generic diagnosis, or to the
establishment of a new genus.

In addition to H. sciurus, two other species of Haemulon were caught
and examined for copepods: 20 specimens of the French grunt, H. flavoli-
-neatum (Desmarest), and one specimen of the smallmouth grunt, H. chrys-
argyreum Gunther; none of these specimens were found to have any copepod
parasites.

In 1980, Cressey and Cressey misspelled Ceratocolax as *‘Ceratacolax”’
when redescribing C. euthynni, and throughout the manuscript; we regret
this error.

Acknowledgments

I would like to thank Dr. Robert Karl Johnson of the Field Museum,
Chicago for the fish identifications; Mr. Walter Brown of the Smithsonian
Institution for the scanning electron micrographs; Dr. Roger Cressey,
Smithsonian, for patient advice and assistance; and Dr. Thomas E. Bow-
man, Smithsonian, for reading and commenting on the manuscript. This
paper is contribution number 68 of the Smithsonian Institution’s Investi-
gations of Marine Shallow-Water Ecosystems Program, Reef and Mangrove
Study—Belize, supported in part by the Exxon Corporation. Special thanks
to Dr. Klaus Rutzler, of the Smithsonian, for bringing this program to my
attention, and for many kindnesses.

524 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Literature Cited

Cressey, R., and H. B. Cressey. 1980. Parasitic copepods of mackerel- and tuna-like fishes
(Scombridae) of the world.—Smithsonian Contributions to Zoology 311: iv + 186 pages,
figures 1-139.

Vervoort, W. 1965. Three new species of Bomolochidae (Copepoda, Cyclopoida) from tropical
Atlantic tunnies.—Zoologische Verhandelingen 76:3—40, figures 1-22.

Department of Invertebrate Zoology, Smithsonian Institution, Washing-
ton, D.C.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 525-534

A REDESCRIPTION OF OCTOPUS ORNATUS GOULD,
1852 (OCTOPODA: CEPHALOPODA) AND THE
STATUS OF CALLISTOCTOPUS TAKI, 1964

Gilbert L. Voss

Abstract.—Octopus ornatus is redescribed based on specimens from Ha-
wail, Bikini Atoll, and Kenya. Variations of mantle shape and color patterns
are described and illustrated as well as internal anatomical features. Callis-
toctopus arakawai is placed in the synonymy of Octopus ornatus. Taki’s
statement that his specimen exhibited luminescence is refuted by observa-
tions of living specimens in Hawaii. As the original type-material is lost, a
large male specimen from Oahu, Hawaii, the type-locality, is designated the
neotype.

Among the cephalopods collected from the R/V Anton Bruun during
United States participation in the International Indian Ocean Expedition
was a series of octopus from East Africa having long dorsal arms and con-
spicuous buff-colored stripes and spots. These animals were at first thought
to belong to Octopus macropus Risso, 1826, but were later considered to
be conspecific with Callistoctopus arakawai Taki, 1964, from Japanese
waters. Examination of material in the United States National Museum of
Natural History disclosed specimens of Octopus ornatus Gould, 1852, that
appeared to be the same. This latter material was received on loan along
with other specimens from Oceania. Finally, two specimens were received
from Hawaii along with color notes made when they were alive. Study of
all of this material leads to the conclusion that Octopus ornatus is a widely
distributed species of which Callistoctopus arakawai Taki is a synonym.
The evidence is presented below.

The measurements and indices used are those of Voss (1963, p. 11) except
for the designation HAI which is the length of the hectocotylized arm as a
percentage of its fellow arm on the opposite side.

Octopus ornatus Gould, 1852
Figs. 1-3

Octopus ornatus Gould, 1852:476, fig. 590, 590a.—Tryon, 1879:112, pl. 30,
figs. 29-30.—Robson, 1929:108.

Polypus ornatus.—Berry, 1909:418; 1914:294, pl. XLVI, figs. 1-2.

Callistoctopus arakawai Taki, 1964:292, pls. 2-3, text-figs. 34-41.

Octopus arakawai.—Dong, 1979:72, pl. 1, fig. 2.

526 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Octopus ornatus: a, Dorsal view of neotype (88 mm ML); b, Color pattern and
shape of 101 mm ML male from East Africa; c, Same of 47 mm ML male from East Africa;
d, Same of 66 mm ML male from East Africa; e, Funnel organ of 101 mm ML male from East
Africa; f, Funnel organ of 74 mm ML female from Hawaii.

VOLUME 94, NUMBER 2 S27

Material examined.—Neotype, male, mantle length 88 mm, from Black
Point, Oahu, Hawaii, collected by Stephen Kempf, 11 January 1976, USNM
730020.—1 female, mantle length 74 mm, Oahu, Hawaii, collected by C. F.
E. Roper and R. E. Young, UMML 1756.—1 male, mantle length 83 mm,
Market, Hawaii (Albatross) S.S.B. #382, USNM 214609.—1 male, mantle
length 98 mm, Enyu Island, entrance to lagoon, Bikini Atoll, Marshall
Islands, Schultz leg. 16 March 1946, USNM 574184.—4 females, mantle
lengths 22-65 mm, USNM 730022.—1 male, mantle length 66 mm, USNM
730023, R/V Anton Bruun, cruise 9, sta. HA-1, Andromache Reef just south
of entrance to Port Kalindi, Mombasa Harbor, Kenya in 0-1 meter, 15
November 1964.—2 males, mantle lengths 47-101 mm, same data as above,
UMML 1756.—3 males, mantle lengths 44-104 mm, R/V Anton Bruun,
cruise 9, sta. HA-2, Andromache Reef, just south of entrarfce to Port Kal-
indi, Mombasa Harbor, Kenya, in 0-1 meter, 16 November 1964, USNM
730021.

Description.—The description is based upon 15 specimens ranging in dis-
tribution from Hawaii to Bikini Atoll and East Africa. While the characters
are consistent and show little variation over the species’ range, some dif-
ficulties arise from the ability of the animals to elongate or contract the
mantle. Round and tubular shaped mantles are found in the material and
can have drastic effects upon indices. In the elongate stage the mantle length
is greatly increased resulting in low indices in some characters and high
ones in others. This problem should be borne in mind when using the in-
dices.

The description is based primarily upon the large male from Oahu des-
ignated as the neotype.

The mantle is round, bulbular, smooth posteriorly or with a small terminal
point, or it may be greatly elongate and tubular (MWI—males, 31-62./-83;
females, 27-60.7-70). It is muscular and thick-walled with a moderately
wide aperture. The funnel is muscular, stout, and free for about half of its
length. The funnel organ is W-shaped with the outer limbs a little shorter
than the median ones.

The head is narrower than the mantle with (HWI—males, 32-45.9-S7;
females, 31-46.2—55) and in the specimens with round mantles there is a
distinct neck region. In those with tubular mantles no neck region is found.
The eyes are of moderate size and are somewhat protuberant.

The arms are long and stout (MAI—males, 15-—/8.2-—23; females, 14—
18.8—26; ALI—males, 82-84.8—87; females, 80-84.5-88), but taper to slen-
der tips. The arm order in nearly all cases is I.II.IHI.IV with I the longest
(two exceptions). The suckers are biserial and large (SIn—males, 7.9-//.4—
15; females, 5—/0—14) and there are no specially enlarged suckers in the
males. The web is shallow (WDI—males, 9-//.4-—14; females, 8—/0.3-12).

528 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Octopus ornatus: a, Hectocotylus of 47 mm ML male from East Africa; b, Hec-
tocotylus of neotype (88 mm ML); c, Hectocotylus of 98 mm ML male from Bikini; d, Hec-
tocotylus of 101 mm ML male from East Africa; e, Lower beak, f, Upper beak, of 101 mm
ML male from East Africa.

The web formula is very variable but sector B is usually the deepest and
sector E is usually the shallowest.

The third right arm is hectocotylized and is shorter than its fellow arm
(HAI 59-68.7-82) (Fig. 2a—d). There is a distinct well-formed spermato-
phoral groove. The ligula is small (LLI—2.3—5.4-6.9) and is elongate with
strongly inrolled lateral margins completely enclosing the deep, smooth lon-
gitudinal groove on the oral face. In small specimens the ligula is more open
while in the specimen from Bikini the distal fifth of the ligula margins is

VOLUME 94, NUMBER 2 29

Fig. 3. Octopus ornatus: a, Digestive tract from 101 mm ML male from East Africa; b,
Digestive tract from 74 mm ML female from Hawaii; c, Genitalia from 98 mm ML male from
Bikini; d. Genitalia from 101 mm ML male from East Africa; e, Radula from 101 mm ML male
from East Africa; f, Radula from 83 mm ML male from Hawaii.

fused together as shown in figure 2c. The calamus is small (CLI—13.6—23.7-
39) and set flush with the margins of the ligula.

There are 12—14, more often 14, lamellae on the outer demibranch of the
gills.

Several specimens were dissected to permit description of the digestive
and reproductive tracts. The beaks are distinctive. The upper beak has a
well-marked long notch in the cutting edge while the lower beak has a sharp
ridge medially in the lamella which is divided in the posterior half. The

Table 1.—Indices and other data of 9 males of Octopus ornatus.

Index Africa Africa Africa Hawaii Hawaii Africa Bikini Africa Africa
Mantle length 44 47 66 83 88 98* 98 101* 104*
Mantle width I 68 83 V7 67 78 45 iI 33 31
Head width I 50 53 53 42 57 35 56 32 35
Mantle arm I 15 16 17 15 16 22 18 2D, 23
Arm length I 87 86 86 87 86 82 85 82 82
Arm width I nS) OD 23 7/ 24 18 — 17. 13
Web depth I 11 11 11 12 9 14 — 12 oo
Sucker Index

normal 13 13 13 10 15 10 12 9 7.9
Hect. arm I 67 73 82 59 Ww 60 — 61 —
Ligula length I 2.8 2.3 6.9 4.6 6.6 4.9 6.0 6.2 —
Calamus length I 39 30 13.6 D5 18.8 Bell — 16.7 —
Arm formula 253-4 120324) 12384) 1243 24531234 eer lena,
Web formula BDCAE BACDE BCADE ACBDE BACDE ACBDE DBCEA BACDE_ ?
Gills 12 13 13 13 14 14 14 13 14

* indicates elongate forms.

radula from both Hawaiian and East African specimens are very similar.
Both have a symmetrical seriation of A®&° with an average of A’, with about
2 cusps on each side of the rachidian. The digestive tract is very similar
between the Hawaiian and East African specimens. Both show elongate
posterior salivary glands, an enormous crop, large stomach and separated
Spiral caecum. The digestive gland of the Hawaiian female is elongate in the
anterior part but wide posteriorly and has a 2-parted ink sac, the largest
posterior. The East African specimen has an elongate gland (it is from a
specimen with a tubular mantle) and a small posterior ink sac. Both glands

Table 2.—Indices and other data of 6 females of Octopus ornatus.

Index Africa Africa Africa Africa Africa Hawaii
Mantle length WD) 33 47 64* 65 74
Mantle width I 68 70 66 Dy 63 70
Head width I 55) 48 43 31 45 SS»)
Mantle arm I 14 18 16 26 18 Dil
Arm length I 88 85 86 80 85 83
Arm width I 20 19 14 9 17 32
Web depth I 8 10 10 10 12 12
Sucker Index normal 9 14 11 5 9 12
Arm formula 1.2.3.4 1.2.3.4 1.2.3.4 ORG, MpWBL! 2.1.3.4
Web formula ABCDE BACDE BCDAE A=CBDE BCDA=E BCD=AE
Gills 14 14 14 14 14 12

* indicates elongate forms.

VOLUME 94, NUMBER 2 Sail

are obviously modified for elongation. The posterior end of the glands is
strongly bilobed.

The female genitalia are not illustrated as none of the specimens with
viscera in good condition were adult. However, the ovary is not posterior
but median and ventral in position, oval or squarish, with very long slender
proximal and distal oviducts with small oviducal glands.

The male genitalia are very similar in all specimens. The one from Bikini
has a long penis but over half of it is composed of a posterior diverticulum
that is folded under itself and projects forward. The penial apparatus from
an East African specimen conforms to this same general pattern but is longer
and more slender, coming from an elongate-mantled specimen. The sper-
matophores of all the males were too incomplete to measure or illustrate.

The skin of the mantle, head and aboral surface of the arms and web is
finely to roughly granulose, resembling that of Octopus vulgaris in some
forms. There are a few large contracted papillae over each eye but not large
enough to be classed as ocular cirri.

The color of the more recently preserved material at my disposal is pur-
plish red dorsally on the mantle, head and arms. In long-preserved speci-
mens the background color is reddish brown. Ventrally and on the oral
surface of the arms and web the skin is a pale flesh color. The pattern of
splotches, spots and stripes is very similar in all of the specimens and is
well shown in the figures. Variations of mantle shape and color pattern are
shown in the outline drawings. These spots and stripes are pale to bright
buff. In some specimens, particularly those from East Africa, the spots and
stripes have a raised ridge or crest or fleshy mound in their centers, the
latter, especially on the arms, reminiscent of those found in Octopus mac-
ropus.

Type.—The original type-material has been lost. The neotype, a male,
mantle length 88 mm is USNM 730020.

Type-locality.—Sandwich Islands, collected by the U.S. Exploring Ex-
pedition.

Discussion

The original specimens were from the Wilkes expedition “‘obtained at the
Sandwich Islands at Oahu and Mauii.’’ None of these specimens have been
located in any of the museums where they might be expected to have been
deposited, and all of these specimens in alcohol must be assumed to be lost
(personal communication, Richard I. Johnson, Museum of Comparative Zo-
ology, Harvard). The most important part of the original description is the
following (Gould, 1852, p. 477): ‘““The ground color is deep orange; beneath
somewhat clouded with white; above variegated with five longitudinal buff
Stripes, the median one extending to between the eyes, the two lateral ones
curving on each side, like median lines, and extending only to the neck;

532 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

between these lines, around the middle of the sac, are deep brown patches,
and also between the bases of the arms; there are also brown mottlings
along the back of the arms. These, with the pale, bubble-like patches around
the base of the sac and along the arms, give a very gay and diversified
coloring.’

This description of the color pattern almost exactly suffices for a descrip-
tion of the colors of the specimens from East Africa and of Taki’s Callis-
toctopus arakawai (Taki, 1964: pl. 2).

An apparent discrepancy is the extremely narrow and elongate body of
the African specimens. Some of these have narrower mantles than are
known for adults of any other species of octopod. This is variable, however,
and several specimens of normal mantle shape are in the series.

Examination of the digestive and reproductive tracts of both normal and
elongated specimens shows that this species is especially adapted for shape
change. This is seen in the elongated digestive gland (Taki, 1964, p. 295,
noted that it nearly filled the mantle), the very long ink sac duct, and the
long convoluted oviducts and anteriorly located ovary.

The two radulas figured, one from Bikini and the other from East Africa,
are nearly exactly the same with symmetrically placed cusps on the rachid-
ian and in a series of 7 to 8. Taki (1964, p. 294, figure 39) also shows a
symmetrical series of 7 rather than 6 as he states in the text.

A variable feature requiring further study is the hectocotylus. Unfortu-
nately, the hectocotylus described and figured by Berry (1914, pl. XLVI,
fig. 2) is from an immature specimen and resembles that of immature spec-
imens in the present material. The single male from Hawaii available to me
has a long, slender but fully excavated ligula. The male from Bikini collected
by Schultz has a narrow hectocotylus with a strongly inrolled ligula whose
margins are fused together along the terminal fifth of its length. The question
arises: does this represent an abnormality or a geographic variation? The
African specimens are very similar to Taki’s specimens from Japan and may
represent the more ‘‘typical’’ condition. ©

Taki erected the genus Callistoctopus for his new species arakawai and
magnocellatus. The characters by which he differentiated the genus from
Octopus are ‘‘(1) large size, (2) robust construction, (3) characteristic color
pattern, (4) reduction of ink-sac.’’ The size and robustness of construction
are no greater than in many other species of Octopus, nor is the reduction
in size of the ink sac of particular note. All of the specimens described here
have a distinctive color pattern, in some ways resembling that of Octopus
macropus (i.e. pale spots on body and arms). Certainly magnocellatus
should not have been included as its color pattern is very dissimilar to that
of arakawai and should be included among the regular ocellated species of
Octopus. There is no valid basis for the retention of the genus Callistoc-
topus and it is placed in the synonymy of Octopus. C. arakawai is consid-

VOLUME 94, NUMBER 2 555

ered to be a synonym of O. ornatus Gould, 1852, a widely distributed Indo-
West Pacific species.

Dong (1979: 72) has reported the occurrence of arakawai from off Xisha,
Guandong Province, China, placing it correctly in the genus Octopus. A
photograph of the specimen (pl. 1, fig. 2) shows that it is a long-bodied
specimen resembling the elongate animals from East Africa.

A final question needs to be examined. Taki (1964:296-298, 301-304) stat-
ed that the living animal, although not personally seen by him, emitted
‘bluish phosphorescent light when stimulated; the light is emitted from the
pink stripes and spots which are scattered all over the body surface except
the ventral surface of body and arms; at that time the colour pattern looks
thin-coloured than ordinary time.’’ Taki’s examination of the cellular struc-
ture of these “‘phosphorescent’’ areas revealed deeply-buried cells that he
interpreted to be light producing cells but that seemed to me to be iridocytes
or reflecting cells, especially as the latter are not mentioned in his histolog-
ical comments. To verify this, I requested that Drs. Roper and Young, both
with extensive knowledge of cephalopod luminescence, examine living spec-
imens in Hawaii, if such could be found.

In a letter from Roper dated 20 November 1975 he wrote ‘‘The so-called
luminescence surely is not true luminescence. Dick (R. E. Young) and I
Saw it and the girl (graduate student using the specimen for behavioral stud-
les) saw it repeatedly under fairly well lit conditions. The animal pulls back
its chromatophores and exposes the long patches of iridophores which ex-
hibit iridescence and shimmer. It is vivid enough to be mistaken for lumi-
nescence—if you didn’t look too carefully.’’ The statement ‘‘pulls back its
chromatophores”’ corresponds to Taki’s “‘the colour pattern looks thin-col-
oured than ordinary time.”’

Roper and Young’s observations amply refute the existence of biolu-
minescence in this species and to date no octopod is known to possess
luminescent organs (Thomas, 1977, p. 386, has shown that the supposed
light organs in Tremoctopus lucifer Akimushkin, 1963, are not light organs).
The specimen from which Roper and Young’s observations were made died
shortly thereafter and was sent to me. It is a female and corresponds to the
others in all particulars.

There is a complex of species in the Indo-West Pacific region character-
ized by the possession of long arms I. These animals have generally been
assigned, without critical study, to Octopus macropus. In the light of the
situation with regards to Octopus ornatus, other members of this complex
need reexamination to determine their specific relationships.

Acknowledgments

I wish to thank Clyde F. E. Roper of the United States National Museum
of Natural History for the loan of material from that institution, for reading

534 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

the manuscript, and both Roper and Richard E. Young, University of Ha-
wail, for kindly producing specimens from Hawaii and for providing notes
on the colors of the living animal. Michael Sweeney of the USNM kindly
assisted me on a visit to Washington and gave unsparingly of his time. The
illustrations of color patterns and ligulas are the work of Banjong Mian-
manus, the full view of the neotype is by Laura Sartucci. The others are
my own. I also wish to thank the National Science Foundation for its con-
tinued support of my research by Grant DEB-7603456 for which this is a
report. This paper constitutes a scientific contribution from the Rosenstiel
School of Marine and Atmospheric Science, University of Miami.

Literature Cited

Akimushkin, I. I. 1963. Cephalopods of the Seas of the U.S.S.R. (1965 English translation of
Golovonogie mollyuski morei SSSR).—Izdatil’stvo Akademii Nauk SSSR, Moskva,
223 pp.

Berry, S. Stillman. 1909. Diagnoses of new cephalopods from the Hawaiian Islands.—Proc.

U.S. Natl. Mus. 37:407-419, text-figs. 1-9.

. 1914. The Cephalopoda of the Hawaiian Islands.—Fish. Document No. 789:257-362,

11 pls., 40 text-figs.

Dong, Zhengzhi. 1979. A preliminary report of the cephalopods from the Xisha waters, Guang-
dong Province, China.—Studia Marina Sinica, no. 15:71-74, pl. 1.

Gould, Augustus A. 1852. United States Exploring Expedition etc., under the command of
Charles Wilkes. Volume 12, Mollusca and shells. 1-510 pp., atlas of 52 pls. Philadelphia.

Robson, G. C. 1929. A monograph of the recent Cephalopoda. Part I. Octopodinae.—Bnitish
Museum (Natural History) London, pp. 1—236, 7 pls., 89 text-figures.

Taki, Iwao. 1964. On eleven new species of the Cephalopoda from Japan, including two new
genera of Octopodinae.—J. Fac. Fish. Animal Husb., Hiroshima Univ. 5(2):277—330, 7
pls., 68 text-figs.

Thomas, Ronald F. 1977. Systematics, distribution, and biology of cephalopods of the genus
Tremoctopus (Octopoda: Tremoctopodidae).—Bulletin of Marine Science 27(3):353-
3977

Tryon, George W. 1879. Manual of conchology. Vol. i, Cephalopoda. Philadelphia, 316 pp.,
112 pls.

Voss, Gilbert L. 1963. Cephalopods of the Philippine Islands. U.S. Natl. Mus. Bull. 234, 180
pp., 4 pls., 36 text-figs.

Rosenstiel School of Marine and Atmospheric Science, University of
Miami, 4600 Rickenbacker Causeway, Miami, Florida 33149.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 535-560

A REVISION OF THE NETTASTOMATID EEL GENERA
NETTASTOMA AND NETTENCHELYS (PISCES:
ANGUILLIFORMES), WITH DESCRIPTIONS
OF SIX NEW SPECIES

David G. Smith, James E. Bohlke, and P. H. J. Castle

Abstract.—The genera Nettastoma Rafinesque and Nettenchelys Alcock
form an easily recognized group within the family Nettastomatidae. The
posterior nostril is displaced dorsally to the top of the head or behind it.
The body is deepest in the middle and tapers at both ends. Nettastoma has
a longer snout and 11-14 pores along the upper jaw; Nettenchelys has a
shorter snout and 8-10 pores along the upper jaw. The position of the pos-
terior nostril varies greatly among species of the two genera. The larva in
both genera is deep-bodied anteriorly and tapers sharply toward the tail.
Nettastoma contains four species. Nettastoma melanura Rafinesque (At-
lantic), N. syntresis n. sp. Smith and Bohlke (Atlantic), and N. solitarium
n. sp. Castle and Smith (Indo-Pacific) have the posterior nostril above the
anterior edge of the eye. They are distinguished inter alia by the number
of supraorbital and lateral-line pores and the relative size of the median
vomerine teeth. Nettastoma parviceps Ginther has the posterior nostril
over the posterior edge of the eye. Metopomycter denticulatus Gilbert is
synonymized with Nettastoma parviceps, and the genus Metopomycter is
Synonymized with Nettastoma. Nettenchelys contains five species. Net-
tenchelys pygmaeus n. sp. Smith and Bohlke (Atlantic) and N. gephyra n.
sp. Castle and Smith (Indo-Pacific) have an enlarged median series of an-
terior vomerine teeth, and the posterior nostril is slightly behind the pos-
terior edge of the eye; N. pygmaeus is mature at 150-200 mm TL, N.
gephyra at about 400 mm. Nettenchelys inion n. sp. Smith and Bohlke,
(Atlantic), N. exorius n. sp. B6hlke and Smith (Atlantic), and N. taylori
Alcock (Indo-Pacific) lack the enlarged anterior vomerine teeth and have
the posterior nostril at or behind the occiput. In N. inion the posterior
nostril is just in front of the supratemporal canal, in N. exorius slightly
behind the occiput, and in N. taylori between the dorsal-fin origin and the
anus.

The Nettastomatidae are a rather poorly known family of eels inhabiting
the outer continental shelf and the continental slope of the world’s tropical
and warm temperate oceans. Closely related to the Congridae, nettasto-
matids are characterized primarily by an elongation and narrowing of the

536 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

head and snout and an enlargement of the mouth. The teeth are generally
small and in multiserial bands. The body is elongate and the tip of the tail
is greatly attenuated. Pectoral fins are usually absent. The distinction be-
tween the Congridae and the Nettastomatidae is not clear, and further study
may lead to the union of the two families.

One readily distinguishable group within the Nettastomatidae is composed
of the nominal genera Nettastoma Rafinesque, Metopomycter Gilbert, and
Nettenchelys Alcock. In this group the posterior nostril is displaced dorsally
to the top of the head or even farther back in some species. The body is
deepest in the middle and tapers toward both ends. The larva in these forms
is deep bodied anteriorly and narrows sharply toward the tail (Smith
1979:11).

This group was known for many years from a single Atlantic species of
Nettastoma, N. melanura Rafinesque, and two poorly defined Pacific
species of Metopomycter, M. parviceps (Gunther) and M. denticulatus Gil-
bert. These two genera were distinguished by the position of the posterior
nostril, near the anterior margin of the eye in Nettastoma and near the
posterior margin of the eye in Metopomycter. Nettenchelys was so poorly
known that no attempt had been made to determine its relationships; its
omission from Jordan’s Genera of Fishes (Jordan 1917—1920) may have con-
tributed to its obscurity.

We have recently discovered several additional species of this group in
both the Atlantic and Indo-Pacific. The holotype and only known specimen
of Nettenchelys taylori Alcock was reexamined and determined to belong
to the group. The position of the posterior nostril is now seen to form a
graded series among the species, and the basic distinction between Netta-
stoma and Metopomycter has largely disappeared. However, a character
that does divide these species clearly into two groups is the length of the
snout and the number of pores along the upper jaw. Nettastoma melanura,
N. syntresis n. sp., N. solitarium n. sp. and N. parviceps have 11 to 14
pores in this series and a correspondingly longer snout. The remaining
species have a shorter snout and eight to ten pores. We recognize these two
groups as genera, Nettastoma and Nettenchelys. Metopomycter becomes
a synonym of Nettastoma.

Methods

Head length is measured to the upper corner of the gill opening. The
upper jaw is measured from the tip of the snout to the posterior end of the
maxilla. Interbranchial distance is measured between the lower end of the
gill openings across the ventral midline. Depth is measured at the anus.
Head pores are designated as in Smith and Kanazawa (1977) except that
there is no postrictal pore in the infraorbital canal; infraorbital pores are

VOLUME 94, NUMBER 2 S3i7/

MARA A Ana Aa, SANA VN Hr
a he UU AAAS MACANAD AANA AA

at figs
ees Og
Cadeeereeey Ss tie ee

D>
4qaa%

Fig. 1. Diagrammatic view of head of a nettastomatid to show sensory canals and pores.
Abbreviations explained in text.

given as those along the upper jaw plus those behind the eye. The presence
or absence of the fifth supraorbital pore (SO,) is diagnostic in the species
of Nettastoma, and its position is indicated in Fig. 1. The anteriormost
lateral-line pore is located at the junction of the lateral-line and the supra-
temporal and preoperculomandibular canals. We call this a lateral-line pore
throughout, although in the holotype of Nettenchelys taylori it is located
below the level of the lateral-line canal. Lateral-line pores counted are those
anterior to the anus. In the descriptions, the number of Specimens that each
count or measurement was based upon is given in parentheses following
that value. Unless otherwise stated, lengths are total lengths (TL).

Abbreviations for pores are as follows: LL = lateral-line, POM = pre-
operculomandibular, IO = infraorbital, SO = supraorbital, ST = supra-
temporal.

Abbreviations for museums are as follows: AM, Australian Museum, Syd-
ney; ANSP, Academy of Natural Sciences of Philadelphia; BMNH, British
Museum (Natural History), London; BPBM, Bernice P. Bishop Museum,
Honolulu; MB, Museu Bocage, Lisbon; MNHN, Muséum National
d'Histoire Naturelle, Paris; RUSI, Rhodes University, Grahamstown, South
Africa; SAM, South African Museum, Capetown; TM, Tasmanian Museum,
Hobart; UMML, Rosenstiel School of Marine and Atmospheric Science,
University of Miami, Florida; USAIC, Ichthyological Collection, University
of South Alabama, Mobile; USNM, National Museum of Natural History,
Washington, D.C.; ZMUC, Zoological Museum, University of Copenhagen;
ZSI, Zoological Survey of India, Calcutta.

Genus Nettastoma Rafinesque, 1810

Nettastoma Rafinesque, 1810:66 (type-species Nettastoma melanura Rafi-
nesque, 1810, by monotypy).

538 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

@ N. melanura O N.syntresis ® N. solitarium @ N. parviceps

Fig. 2. Distribution of the species of Nettastoma.

Hyoprorus Kolliker, 1854:101 (type-species Hyoprorus messinensis Kolli-
ker, 1854, by monotypy).

Metopomycter Gilbert, 1905:585 (type-species Metopomycter denticulatus
Gilbert, 1905, by original designation).

Muraenosaurus Osorio, 1909:14 (type-species Muraenosaurus Giintheri
Osorio, 1909, by monotypy). Preoccupied by Muraenosaurus Seeley,
1874, Reptilia.

Osorina Whitley, 1951:68 (type-species Muraenosaurus Giintheri Osorio,
1909, by original designation). To replace Muraenosaurus Osorio.

Description.—Body elongate, round in cross section, deepest in middle
and tapering at both ends. Tail slender and attenuate, tip delicate and easily
damaged. Pectoral fin absent. Dorsal fin originates above gill opening. Head
elongate, slightly depressed, snout length about 35-49% head length; ante-
rior nostrils tubular, opening laterally just behind tip of snout; posterior
nostrils displaced dorsally near anterodorsal or posterodorsal margin of eye;
tip of snout soft, flexible, extending a short distance in front of the premax-
illary teeth. Teeth relatively small, in multiserial bands or patches. Maxillary
and mandibular teeth in bands, innermost teeth somewhat enlarged; pre-
maxillary tooth patch short, continuous with maxillary teeth; vomerine
tooth patch elongate, reaching level of eye, median teeth sometimes en-
larged. Eleven to fourteen infraorbital pores along upper jaw, the second
one slightly elevated in short ascending branch of infraorbital canal imme-
diately behind anterior nostril (Fig. 1); five or six supraorbital pores includ-
ing one near posterior end of canal above eye; third supraorbital pore slight-

VOLUME 94, NUMBER 2 539

Oe Be oe ee
ae NOBORU PERE gaE SET ie eS SesCcn Te Pee eee epee eee.

Fig. 3. Nettastoma parviceps, AM 1.20452-003 (634 mm incomplete). A, Head. B, Upper
dentition. C, Lower dentition. D, Dorsal view of a portion of head to show posterior nostrils
and posteriormost supraorbital pore. Scales in mm; a single scale applies to B and C.

ly posterior to second infraorbital pore (i.e., the one in ascending branch of
infraorbital canal); three supratemporal pores. Viscera: stomach pale, reach-
ing near level of vent; intestine pale. Maximum TL probably 800-1000 mm.
Range.—Eastern and western tropical Atlantic, including Mediterranean;
Indo-Pacific from East Africa to Hawaii, Australia to Japan (Fig. 2).
Etymology.—Gr. netta, duck; Gr. stoma, mouth. The gender is neuter.
Note.—Nettastoma elongatum Kotthaus, 1968 is a species of Saurenche-
lys Peters.

Key to the Species of Nettastoma

1. Posterior nostril near posterior margin of eye (Indo-Pacific) .......
2p 0. o-alcsshath eiucatec eet t Me R l eo R OOS nal tlagran N. parviceps Gunther

= SEOSteHOr NOSthl nearn antenor margin Of Cye ..........-.--.-...- y
2. SO; pore usually absent; median vomerine teeth slightly enlarged;

ISIE pores 44=49) (Atlantic)... 02) «2s tee a N. melanura Rafinesque
SO pOTreusuallyspresemt. Il ponesi4oxOmlewer 57... 2 oo. 3
3. LL pores 38-41; median vomerine teeth not enlarged (western

FNUIAINUIC) ere e aes Meee ak, Se eee ee nes: Me eee N. syntresis n. sp.

— LL pores 43-45; median vomerine teeth somewhat enlarged (Indo-
ACHIC) eee ee ee eRe Ee eee Re hs eeowice N. solitarium n. sp.

540 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Nettastoma parviceps Gunther, 1877
Bigs ele

Nettastoma parviceps Gunther, 1877:446 (original description, Yedo, Japan,
holotype BMNH 1879.5.14.440).

Metopomycter denticulatus Gilbert, 1905:585, fig. 233 (original description,
Hawaii, holotype USNM 52191).

Metopomycter parviceps: Gilbert, 1905:585.

Diagnosis.—SO; pore normally absent (Fig. 3A). LL pores 49-58. Median
vomerine teeth not enlarged (Fig. 3B). Posterior nostril above posterodorsal
corner of eye (Fig. 3D).

This species has appreciably more lateral-line pores than N. melanura,
N. syntresis, and N. solitarium, and differs from these species in having the
posterior nostril over the posterior margin of the eye rather than over its
anterior corner.

Description.—Meristic characters: Total vertebrae unknown but at least
230. Pores: LL 49-58 (33); POM 15-20 (23), IO 11-14 (24) + 3 (29); SO 5-6
(24); ST 3 (23; one specimen had 4).

Morphometric characters: Predorsal length 25-32% preanal length (37);
head 27-34% preanal (37); snout 36—49% head (36); eye 9-14% head (37);
upper jaw 50-66% head (36); gill opening 3—9% head (37); interbranchial 7—
17% head (35); depth 4—-10% preanal (32).

Dentition (Fig. 3B, C): Maxillary and mandibular teeth in bands, the inner
teeth slightly larger; premaxillary tooth patch short, continuous with max-
illary teeth; vomerine tooth patch elongate, reaching level of eye, median
teeth not enlarged.

Color: A freshly caught specimen (AM I.20452-003) was gray; vertical
fins black-edged; snout gray-black; iris and opercular regions silver.

Size: The largest specimen examined was 815 mm TL, 335 mm preanal.
Females with large eggs were about 500-600 mm TL.

Variation.—Nettastoma parviceps is known from four widely separated
areas: Japan, Hawaii, Australia, and South Africa. These four populations
differ from each other in four characters (Table 1). In addition, the two
northern hemisphere populations differ from the two southern hemisphere
populations in the nature of the posteriormost supraorbital pore (the one
between the posterior nostrils). In the former this pore is paired, whereas
in the latter the pores of the right and left sides are fused into a single
median pore (Fig. 3D).

Specimens of N. parviceps almost invariably have damaged tails, making
it impossible to give accurate vertebral counts. Several specimens in which
the caudal region seemed to be intact or nearly so yielded counts of about
230. The holotype of N. parviceps (from Japan) has approximately 234
vertebrae with 53 preanal; that of Metopomycter denticulatus (from Hawaii)

VOLUME 94, NUMBER 2 54]

Table 1.—Geographic variation in Nettastoma parviceps.

Predorsal length
as percentage

of preanal
length 25 26 27 28 29 30 31 32
Japan 1 0 D, 3 1
Hawaii 3 5 3 4
South Africa 2, 3 3 1 1 2, 1
Australia 1 D

Head length as
percentage of

preanal length 27 28 29 30 31 32 33 34
Japan pp 22 3
Hawaii 1 3 5) 6
South Africa 2 D 4 4 1
Australia 1 1 0 1

Snout length as
percentage of

head length 36 37 38 39 40 41 42 43 44 Aa 49
Japan 3 1 3)
Hawaii 1 3 6 3 1 1
South Africa 2 0 1 3 3 1 l 0 1 1
Australia 1 1 0 0 0 1

Preanal
lateral-line
pores 49 50 51 52 53 54 55 56 57 58

Japan 1 5 0 1
Hawaii 2 4 2 4 2
South Africa 3 1 4 1 0 p
Australia 1 2

has 158 with 50 preanal, but the caudal region is obviously incomplete.
Myomere counts from nine larvae (all southern hemisphere) ranged from
ca. 220 to 236.

Range.—This species (Fig. 2) has been recorded from Japan, Hawaii,
eastern Australia from New South Wales to Tasmania, and southeastern
Africa from southern Mozambique to near the Cape of Good Hope. Depth
60—1,190 m (mean 575 m). It has a bitemperate distribution, being rare or
absent in equatorial waters.

Study material.—JAPAN: BMNH 1879.5.14.440 (holotype of Nettastoma
parviceps: 665), CHALLENGER 232, 35°11'N, 139°28’E, 630 m, 12 May
1875. USNM 112037 (1:546), ALBATROSS D5059, 33°05'30"N, 138°39'50"E,
13 Oct. 1906. USNM 148872 (1:458) and USNM 149779 (1:518), ALBA-
TROSS D4957, 32°36'N, 132°23'’E, 799 m, 23 Aug. 1906. USNM 149786
(1:495), ALBATROSS D4971, 33°23'30’N, 135°34’E, 1,187 m, 30 Aug. 1906.
USNM 149787 (2:495-537), ALBATROSS D4972, 35°25'45"N, 135°33’E, 805

542 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

100° 2 err 90 85° : 80° 75° 70° 65° 60° 55°
Ssitiiohfichotohih foot satanaeeasaninuivasenensivadusenssvadatanisvesetanssuscasestavsoniovesssantavisotentavsinientareaat fob Sratvaenaensineiuesetontansanssntsnsdnssuisvaseinuinusnasvaivsivssersntintsnisoavaiviivssnsatintinisnapraiviivistiaitintins intansiniinterd

@ N. melanura

e) Ss i
a on N. syntresis

ier.
ean

‘ o
race

J

o
CARACAS

~_ Orifiec

Fig. 4. Distribution of Nettastoma melanura and N. syntresis in the western North Atlan-
tic.

m, 30 Aug. 1906. USNM 164157 (1:590), ALBATROSS 5067, 35°05'50’N,
138°41’1S’E, 15 Oct. 1906.

HAWAII: BPBM 24856 (12:487-755), TOWNSEND CROMWELL 61-
66, 21°01'N, 156°08’W, 860 m, 26 Oct. 1972. BPBM 25073 (1:487), TOWN-
SEND CROMWELL 52-87, Hawaiian Islands, Pailolo Channel, mid-chan-
nel, 650 m, 10 March 1971. BPBM 25081 (1:602), TOWNSEND CROM-
WELL 59-20, 21°09’N, 156°11’W, 100-800 m, 11 July 1972. USNM 52191
(holotype of Metopomycter denticulatus: male, 765), Hawaii, 1899.

EASTERN AUSTRALIA: AM 1.19376-002 (1:200 incomplete), KAPA-
LA, 35°44’S, 150°38’E, 550 m, 10 Nov. 1976. AM 1.20452-003 (2:575-—634
incomplete), KAPALA 75-05-05, 33°32’S, 152°00’E, 823 m, 19 Aug. 1975.
AM 1.20452-004 (1:370 incomplete), KAPALA, 33°31’S, 152°00’E, 822 m,
19 Aug. 1975. TM D1649 (1:480), 43°57’S, 150°23’E, 730-940 m, 30 Dec.
1979.

SOUTHEASTERN AFRICA: ANSP 123528 (1:231), 25°29’S, 33°35’E,
450 m, 28 Sept. 1964. RUSI uncatalogued (1:915), off Limpopo River, Mo-
zambique, 485 m, 17 Jan. 1968. SAM-12785 (1), PIETER FAURE, off Cape
St. Blaize, South Africa, 60 m. SAM-12786 (1:256), PIETER FAURE, off
East London, South Africa, 357 m. SAM-12787 (3:137-—297), PIETER
FAURE, off East London, South Africa, 732 m. SAM-12788 (1:259), PIE-
TER FAURE, off East London, South Africa, 549 m. SAM-24372 (2:349
and incomplete), Division of Sea Fisheries, off Durban, South Africa, 406
m 26 July 1964.

VOLUME 94, NUMBER 2 543

Etymology.—L. parvus, little, and L. caput, head; the significance is not
obvious, although the head of this species is slightly smaller than that of N.
melanura, the only other species of the genus known to Gunther.

Nettastoma melanura Rafinesque, 1810
Figs. 2, 4, 5A, 6A

Nettastoma melanura Rafinesque, 1810:66 (original description, Sicily, ho-
lotype lost).

Muraenophis saga Risso, 1810:370 (original description, Nice, Holotype
MNHN B-849).

Murenophis saga: Risso 1826:193.

Muraena saga: Costa 1844:10.

Hyoprorus messinensis KoOlliker, 1854:102 (original description, Messina).
Larva.

Leptocephalus longirostris Kaup, 1856:150 (original description, North At-
lantic). Larva.

Nettastoma melanurum Gunther, 1870:48 (unjustified emendation).

Nettastoma mendax Facciola, 1893:147 (original description, Mediterra-
nean).

Nettastoma saga: Facciola, 1899:31.

Muraenosaurus Giintheri Osorio, 1909:14, pl. 1, Fig. 2—4 (original descrip-
tion, Portugal, holotype MB 2066).

Leptocephalus urosema Lea, 1913:39 (original description, near Azores,
two syntypes at Natural History Museum, Bergen). Larva.

Osorina guentheri: Whitley 1951:68.

Not Nettastoma melanura Rafinesque of Facciola 1893:144, 1899:30, Poll
19534129)

Diagnosis.—Usually five supratemporal pores, SO, absent (Fig. 5A). LL
pores 43-49. Median vomerine teeth somewhat enlarged (Fig. 6A). Posterior
nostril near anterodorsal margin of eye.

Description.—Meristic characters: Total vertebrae ca. 190-210. Bran-
chiostegal rays 9 (1). Pores: LL 43-49 (32); POM 18-21 (27); IO 12-14
(29) + 3 (30); SO 5 (29); 6 (4); ST 3 (28; one specimen had 5).

Morphometric characters: Predorsal 33-40% preanal (31); head 35-42%
preanal (31); snout 35—44% head (30); eye 8-11% head (30); upper jaw 51-
62% head (29); gill opening 2-8% head (26); interbranchial 13-29% head
(17); depth 5—12% head (28).

Dentition (Fig. 6A): Maxillary and mandibular teeth in bands, the inner
teeth largest; premaxillary tooth patch short, continuous with maxillary
teeth; vomerine tooth patch elongate, reaching level of eye, median teeth
usually enlarged somewhat, but not forming a single row.

Osteology: Supraoccipital absent. Upper pharyngeals divided. Epicentral
processes on caudal vertebrae.

544 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. A, Head of Nettastoma melanura, ANSP 117074 (581 mm). Note absence of SO,
pore. B, Head of Nettastoma syntresis, ANSP 114965 (433 mm). Note presence of SO; pore.
Scales in mm.

Color: In preservative, brown or gray; vertical fins black-edged.

Size: The largest specimen examined was 773 mm TL, 303 mm preanal.

Variation.—The frequency of regenerated tails makes it difficult to obtain
complete vertebral counts. The few x-rayed specimens that appeared to be
intact yielded counts of slightly over 200. Myomere counts from 15 larvae
ranged from 186 to 211.

Of 33 specimens examined, 29 lacked the SO; pore. Three specimens had
this pore on one side but not the other. One specimen had the pore on both
sides, but they were not directly opposite each other. The presence of the
SO; pore is an anomalous condition in this species.

Despite its wide range, we find no evidence of geographic variation in
Nettastoma melanura.

Nomenclatural note.—During much of the nineteenth century Nettasto-
ma melanura was confused with the other nettastomatid inhabiting the Med-
iterranean, the species now known as Facciolella physonema (Facciola),
which is characterized by the posterior nostril opening through a slit in the
upper lip. Facciola (1893:147) recognized this and concluded that Rafines-
que’s Nettastoma melanura referred to the species with the labial nostril;
he proposed Nettastoma mendax for the species with the superior nostril.
He later (1899) elaborated on this but decided that the species with the
superior nostril should be known as Nettastoma saga (Risso, 1810). Still
later (Facciola, 1911, 1914) he reversed his opinion and decided that N.
melanura Rafinesque really referred to the species with the superior nostril.

VOLUME 94, NUMBER 2 545

Oa COLA spot Se ne SiC Pes Pie ad, PS 6
Bi : OEY ” SE Boo Bpswn Coser SS

Op ER DOS rae
awe re BA PSS
a se SATE AIS 2 = s
SS 9 9 99 I >
4 s >> >

tek cle 1143)

Fig. 6 A, Upper dentition of Nettastoma melanura, ANSP 117074 (581 mm), showing
enlarged median vomerine teeth. B, Upper dentition of Nettastoma syntresis, ANSP 158936
(482 mm). Note median teeth not enlarged. Scales in mm.

This made N. saga and N. mendax synonyms of N. melanura and left the
species with the labial nostril without a name. Facciola (1911:275) gave the
generic name Nettastomella (preoccupied and later replaced by Facciolella
Whitley) and later (Facciola, 1914:47) the specific name physonema to the
latter. Thus at different times Facciola used the name Nettastoma melanura
to refer to two distinct species of eel.

Range.—Widely distributed on both sides of the Atlantic, from the Med-
iterranean to the Gulf of Guinea and from the northern Gulf of Mexico and
east coast of Florida through the Caribbean to the Guianas (Figs. 2, 4).
Depth 37-1647 m (mean 730 m). Nettastoma melanura appears to be absent
from the Santaren and Nicholas Channels, and the Bahamas. It is replaced
there by Nettastoma syntresis n. sp.

Study material.—GULF OF MEXICO: ANSP 117066 (2:567—660), OR-
EGON 4815, 25°05'N, 96°27’W, 732 m, 12 Apr. 1964. ANSP 117074 (1:581),
OREGON 4076, 28°33'N, 86°27’W, 458 m, 3 Dec. 1962. USNM 178997
(2:616, incomplete), OREGON 4149, 29°12'N, 87°48’W, 732 m, 17 Dec.
1962. USNM 179069 (3:440-773), OREGON 4006, 29°03’N, 88°20'W, 641
m, 23 Oct. 1962. USNM 195898 (2:339-544), OREGON 2822, 28°55'N,
87°49'W, 1647 m, 16 July 1960. USNM 198051 (1:666), OREGON 4580,
29°06'N, 88°06’W, 732 m, 11 Dec. 1963.

STRAITS OF FLORIDA: UMML 13212 (1:606), OREGON 4374,

546 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

24°21'N, 83°36’W, 732 m, 7 Aug. 1963. UMML 13727 (1:373), OREGON
4373, 24°26'N, 83°31’ W, 549 m, 7 Aug. 1963. UMML 15648 (1:473), GERDA
362, 24°10’N, 81°42’W, 631 m, 15 Sept. 1964. USNM 185651 (1:294), SIL-
VER BAY 449, 28°42'N, 79°48'’W, 512-576 m, 11 June 1958. USNM 197341
(1:626), SILVER BAY 3516, 24°24’N, 80°00'W, 732-860 m, 9 Nov. 1961.

CARIBBEAN: ANSP 126445 (1:635), OREGON 11231, 8°56’N, 76°53’W,
622 m, 2 Nov. 1970. USNM 198754 (1:727), OREGON 4902, 9°02.4’N,
76°31.5’W, 732 m, 28 May 1964. USNM 219848 (1:780), OREGON 6703,
16°53'N, 61°53’W, 750-842 m, 21 May 1967.

GUIANAS: ANSP 114844 (1:655), OREGON 10800, 7°20'N, 53°04’W,
692 m, 17 Nov. 1969. USNM_ 158933 (1:482), OREGON 2039, 5°47’N,
52°55'W, 37 m, 11 Nov. 1957. USNM 158934 (1: incomplete), OREGON
2011, 7°46'N, 54°36’ W, 732 m, 7 Nov. 1957. USNM 158935 (1:659), OREGON
2030, 7°10'N, 52°55’'W, 549 m, 10 Nov. 1957. USNM 185639 (1:606), OR-
EGON, off Surinam, Sept. 1958.

GULF OF GUINEA: UMML 15145 (4:469-555), PILLSBURY 74,
4°20'N, 9°26’W, 732 m, 4 June 1964. UMML 16940 (1:224), PILLSBURY
51, 4°S6’N, 5°01’W, 329-494 m, 31 May 1964. UMML 20903 (3:200-420),
PILLSBURY 256, 3°45’N, 8°03’E, 408-485 m, 14 May 1965. USNM 198619
(1:337), GERONIMO 199, 1°26.4’W, 8°24’E, 3 Sept. 1963. USNM 198629
(1:406), GERONIMO 2-247-VIII, 4°38’S, 11°01’E, 9 Sept. 1963. USNM
199537 (1:460), GERONIMO 206, 2°00’S, 8°47’E.

MEDITERRANEAN: USNM 49345 (1:602), Nice, France.

Etymology.—Gr. melan, black, dark, and Gr. oura, tail, referring pre-
sumably to the dark edges of the posterior part of the dorsal and anal fins.
The correct form of the name would seem to be melanura and not the
emended form melanurum, despite the fact that Nettastoma is neuter. The
Greek ‘“‘oura’’ is a feminine noun, and melanura could legitimately stand
as an appositional noun. There is nothing in the original description to in-
dicate whether Rafinesque intended it as an adjective or a noun. The amend-
ed article 30() of the International Code of Zoological Nomenclature states
clearly that in such cases the name is to be treated as a noun in apposition,
hence no justification exists for emending it to melanurum.

Nettastoma syntresis, new species, Smith and Bohlke

Holotype. —USNM 219849 (male: 442), SILVER BAY 2458, Santaren
Channel, 23°40’N, 79°18’W, 531 m, 5 Nov. 1960.

Diagnosis.—Normally six supraorbital pores, SO; present (Fig. 5B). LL
pores 38—41. Median vomerine teeth not enlarged (Fig. 6B). Posterior nostril
near anterodorsal margin of eye.

Description.—Meristic characters: total vertebrae 201 (1). Branchiostegal
rays % on one paratype (USNM 158936). Pores: LL 38-41 (14); POM 18-
19 (5); IO 12-13 + 3 (11); SO 5-7 (16); ST 3 (12).

VOLUME 94, NUMBER 2 547

2 2000
2-0, 0509 294g ,0
fee 0° 8.9008,
x see TS he O Bren ¢ eetee 00202282025
REEL ated Dako oy 7

5° 3°06

Fig. 7. Nettastoma solitarium, holotype (457 mm). A, Head. B, Upper dentition. C, Lower
dentition. D, Dorsal view of a portion of head showing posterior nostrils. Scales in mm; a
single scale applies to B and C.

Morphometric characters: Preanal 35% TL (1); predorsal 37-41% preanal
(13); head 39-43% preanal (13); snout 41-44% head (13); eye 8-11% head
(13); upper jaw 57-63% head (12); gill opening 4—7% head (12); interbran-
chial 10-22% head (9); depth 7-13% preanal (11).

Dentition (Fig. 6B): Maxillary and mandibular teeth in bands, the inner
teeth larger; premaxillary tooth patch short, continuous with maxillary
teeth; vomerine tooth patch elongate, reaching level of eye, median teeth
not enlarged.

Color: In preservative, brown or gray; vertical fins edged in black.

Size: The largest specimen examined was 482 mm TL, 184 mm preanal.

Variation.—Of 16 specimens examined, 12 had six supraorbital pores on
both sides. One specimen had six on one side and five on the other. Three
specimens had six on one side and seven on the other. In no specimen was
the SO, pore absent on both sides.

Range.—This species has been collected only in the Santaren Channel
(between the Cay Sal Bank and the Great Bahama Bank), the Nicholas
Channel (between Cuba and the Cay Sal Bank), and the Northwest Provi-
dence Channel, Bahamas. Depth 458-641 m. Nettastoma syntresis and N.
melanura appear to have disjunct distributions (Figs. 2, 4).

Study material.—Paratypes: ANSP 114965 (1:433), 23°25'’N, 79°40’ W, 458
m, 16 Dec. 1969. UMML 31525 (3:320-409), COLUMBUS ISELIN 137,

548 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

26°07'N, 78°34.1'W, 560-591 m, 27 Sept. 1973. USNM 157939 (1: incom-
plete), OREGON 1341, 22°55'N, 79°16'W, 439 m, 16 July 1955. USNM
158936 (1:482), COMBAT 450, 23°59'N, 79°43'W, 641 m, 24 July 1957.
USNM 197127 (2:270-360), SILVER BAY 2469, 23°59’N, 79°17’ W, 531-549
m, 7 Nov. 1960. USNM 197342 (1: incomplete), SILVER BAY 2475,
24°48'’N, 79°17'W, 549 m, 8 Nov. 1960. USNM 197345 (6:251-453), same
data as holotype. USNM 202953 (1:260), SILVER BAY 3514, 23°20'N,
79°27'W, 549 m, 8 Nov. 1961.

Others (relatively poor condition, not designated as paratypes): MCZ
38780 (1: ca. 370, in two pieces), ATLANTIS 2982A, 22°48'N, 78°50’ W, 384
m, 11 March 1938. MCZ 38781 (1: 122), ATLANTIS 2987A, 23°22’N,
79°56'W, 521 m, 13 March 1938. MCZ 38803 (1: 298), ATLANTIS 2984,
23°16'N, 79°10’'W, 439-458 m, 12 March 1938. MCZ 38811 (1: 179), AT-
LANTIS 3434, 23°10’N, 79°35’W, 476 m, 1 May 1939. MCZ 38814 (1: 250),
ATLANTIS 3438, 23°05.5’N, 79°37'W, 485 m, 2 May 1939.

Etymology.—Gr. syntresis, channel, passage, strait, referring to the San-
taren, Nicholas, and Northwest Providence Channels, the only places the
species has been found. A noun in apposition.

Nettastoma solitarium, new species, Castle and Smith
Figs. 2, 7

Holotype.—MNHN 1979-187 (female: 457), MUSORSTOM Philippines
50, 13°49.2'N, 120°01.8’E to 13°48.2’N, 120°02.5’E, 415-510 m, 25 Mar.
1976.

Diagnosis.—Six supraorbital pores, SO, present (Fig. 7A). LL pores 42—
45. Median vomerine teeth somewhat enlarged (Fig. 7B). Posterior nostril
above anterodorsal corner of eye.

This species is intermediate between N. melanura and N. syntresis in the
number of lateral-line pores. It shares enlarged vomerine teeth with N.
melanura and the presence of the SO, pore with N. syntresis.

Description.—Meristic characters: Number of vertebrae unknown. Pores:
LL 42-45 (3); POM 19 (3); IO 13 + 3 (3); SO 6 (3); ST 3 (3).

Morphometric characters: Predorsal 35-36% preanal (3); head 37-39%
preanal (3); snout 40-41% head (3); eye 9-11% head (3); upper jaw 51-54%
head (3); gill opening 6-8% head (3); interbranchial 14-16% head (3); depth
9-11% preanal (3).

Dentition (Fig. 7B, C): Maxillary and mandibular teeth in bands, the inner
teeth larger; premaxillary tooth patch short, continuous with maxillary
teeth; vomerine tooth patch elongate, reaching level of eye, median teeth
somewhat enlarged, but not forming a single row.

Color: In preservative, brown or gray; opercular region darker; vertical
fins black-edged to wholly black at tip of caudal region.

Size: The largest specimen was 465 mm TL, 179 mm preanal.

VOLUME 94, NUMBER 2 549

Range.—This species (Fig. 2) has been collected off the Philippine Islands
at 415-610 m and off New South Wales, Australia at 502 m, but we know
of it also in the western Indian Ocean (C. Karrer, pers. comm.).

Study material.—Paratypes, PHILIPPINES: MNHN 1979-188 (1:465 in-
complete), MUSORSTOM Philippines 43, 13°50.5’N, 120°28.0’E to 13°
52.3'N, 120°28.6’E, 484-448 m, 24 Mar. 1976. MNHN 1978-189 (1:405 in-
complete), MUSORSTOM Philippines 44, 13°46.9’N, 120°29.5'E to
13°46.2’N, 120°29.8’E, 610-592 m, 24 Mar. 1976.

Paratype, AUSTRALIA: AM 1.20301-016 (1:320 incomplete), KAPALA,
292525, 153°42’'E to 29°55’S, 153°41’E, 502 m, 23 Aug. 1977.

Etymology.—L. solitarius, solitary, alone, lonely, in reference to the iso-
lated and widely scattered distribution of this species in the Indo-Pacific.
An adjective.

Nettenchelys Alcock, 1898

Nettenchelys Alcock, 1898:149 (type-species Nettenchelys taylori Alcock,
1898, by monotypy).

Description.—Body elongate, round in cross-section, deepest in middle
and tapering at both ends. Tail slender and attenuate, tip delicate and easily
damaged. Pectoral fin absent. Dorsal fin originates above or slightly behind
gill opening. Head moderately elongate, slightly depressed; snout length
about 33-37% head length; anterior nostrils tubular, opening laterally just
behind tip of snout; posterior nostrils located on top of head or body behind
posterior margin of eye, degree of posterior displacement varying greatly
among the known species (Fig. 8); tip of snout soft, flexible, extending a
short distance in front of premaxillary teeth. Maxillary and mandibular teeth
in bands, innermost teeth somewhat enlarged; premaxillary tooth patch
Short, continuous with maxillary teeth; vomerine tooth patch elongate,
reaching about to anterior margin of eye, sometimes several enlarged me-
dian teeth anteriorly. Eight or nine infraorbital pores along upper jaw in-
cluding one in short ascending branch of infraorbital canal immediately be-
hind anterior nostril; five supraorbital pores all well anterior to eye; third
supraorbital pore slightly anterior to second infraorbital pore; two or three
supratemporal pores. Viscera: stomach pale, posterior end near level of
vent; intestine pale. Maximum size probably no more than 550 mm TL.

Discussion.—The species within the genus Nettenchelys as defined here
fall into two groups. Nettenchelys pygmaeus n. sp. and N. gephyra n. sp.
have an enlarged median series of teeth at the anterior end of the vomer
(Figs. 10B, 11B). The posterior nostrils in these species are located on top
of the head behind the posterior edge of the eye (Fig. 8A, B). In N. taylori
Alcock, N. exorius n. sp., and N. inion n. sp. the anterior vomerine teeth
are not enlarged (Fig. 12B) and the posterior nostrils are located at or behind

550 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 8. Semidiagrammatic dorsal view of the five species of Nettenchelys showing position
of posterior nostrils. A, N. pygmaeus. B, N. gephyra. C, N. inion. D, N. exorius. E, N.
taylori.

the occipital region (Fig. 8C—E). Nevertheless, these two groups are closer
to each other than they are to the species of Nettastoma, and we feel that
this is best expressed by placing them in one genus.

Range.—Western North Atlantic and tropical Indo-Pacific. All of the
Species are known from very few individuals.

Etymology.—Gr. netta, duck, and Gr. enchelys, eel. The Greek enchelys
can apparently be either masculine or feminine. Alcock did not indicate the
gender in his description, and it cannot be inferred from the name of the
type species. Article 30(a)(1)(2) of the International Code of Zoological No-
menclature dictates that in such circumstances the name is to be considered
masculine.

Key to the Species of Nettenchelys

1. Anterior vomerine teeth forming an enlarged median series; poste-
rior nostrils on top of head slightly behind eye .................. 2

VOLUME 94, NUMBER 2 551

65°
oo

New © N
2S : . pygmaeus
2 » Orleans Ole °
Galveston Qy

N. exorius
Corpus Vl
Christi
4 Mini
iami

Sv

(o}
aracaib aed
Senos

ATH PPT FAT agooee POT TTT TTT COTO POOP TPP TFT PPT TPH jangcogesa PLT DTI TI == {ii}
a sf — === — — a —S o> — [ + SS S|

100° O55 90° 85° 80° 75° 70° 65°

Fig. 9. Distribution of the three species of Nettenchelys in the western North Atlantic.

— Anterior vomerine teeth not enlarged, posterior nostrils at or behind

OCCIPUi Geary: Moats Eos OE oases wet epligon Khs atleat. 3
2. Very small, mature at 150-200 mm TL; posterior nostril just behind
posterior margin of eye (Western Atlantic) ...... N. pygmaeus, Nn. sp.

— Mature at about 400 mm TL; posterior nostril about midway be-
tween posterior margin of eye and occiput (Indo-Pacific) .........

= uci dh sca aS RR acre cere Sat ce ae Re PE ra Re N. gephyra, n. sp.
3. Posterior nostril just in front of supratemporal canal; median supra-

temporal pore present (western Atlantic) ............ N. inion, N. sp.
— Posterior nostril behind supratemporal canal; median supratemporal
POKCHADS EMU ENS. A hele Re ate SR USE Mi ke cdg dal be Gused ee 4
4. Posterior nostril between occiput and dorsal-fin origin (western
AtlantiCn):. 2G a Age Ae of EES GRE oH AA. N. exorius, n. sp.
— Posterior nostril about midway between dorsal-fin origin and anus
dndosBacihic)}) Pye Re AOR ORL yok LR N. taylori Alcock

Nettenchelys pygmaeus, new species, Smith and Bohlke
Figs. 8A, 9, 10

Holotype.—ANSP 142277, formerly part of USAIC 03460 (male: 175),
Gulf of Mexico, 29°52'’N, 87°06.5’W, 128 m, 28 Aug. 1976.

Diagnosis.—Anterior median vomerine teeth enlarged (Fig. 10B). Poste-
rior nostril on top of head just behind posterior margin of eye (Fig. 8A).
Mature at 150-200 mm TL.

552 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

PAT, -

paw? , ses ait
Ty 2 ar Gece
pe 25 = a
es 4 CAE cs Seine
: 7 , OL, ge PE
g ‘ An S * Ae
BO SIEBER Terai t pipe
Peeper)
3

aa =a = Sane
os 7 i 2
We ys SI? 2 DD 7e8e o*

wale

dentition. C, Lower dentition. Scales in mm; a single scale applies to B and C.

Description.—Meristic characters: Pores: LL ca. 40—42 (2); IO 8 + 3 (3);
SO 5 (3); ST 3 (3). Number of vertebrae unknown.

Morphometric characters: Preanal ca. 35-36% TL (4); predorsal 33-35%
preanal (4); head 30-33% preanal (4); snout 34-37% head (4); eye 8-10%
head (4); upper jaw 53% head (2); gill opening 6—7% head (3); depth 7-8%
preanal (4).

Dentition (Fig. 10B, C): Maxillary teeth in bands, teeth of innermost series
somewhat enlarged. Mandibular teeth in bands, those of innermost series
markedly enlarged. Premaxillary tooth patch short, continuous with max-
illary teeth. Vomerine tooth patch elongate, reaching about to level of an-
terior margin of eye, anteriormost part consisting of a single series of en-
larged teeth medially.

VOLUME 94, NUMBER 2 553

Color: In preservative brown, darker above than below. One specimen
had a distinct bicolored appearance, dark above and pale below.

Size: This is a dwarf species. The largest specimen measured only 201
mm TL, 71 mm preanal. The smallest specimen, a female 151 mm TL, 54
mm preanal was filled with large eggs. Because of the small size, we were
unable to obtain satisfactory radiographs or to count the brachiostegal rays
and the POM pores.

Relationships.—This species is closest to N. gephyra of the Indo-Pacific,
and the differences are discussed under that species.

Range.—Two of the four known specimens came from off western Ven-
ezuela, the other two from the northern Gulf of Mexico (Fig. 9). This sug-
gests that the species is probably fairly widely distributed in the western
Atlantic. Its small size and relatively deepwater habitat may make it difficult
to collect. It would easily slip through the mesh of most nets. Other small
eels usually are collected by divers using rotenone, and Nettenchelys pyg-
maeus lives too deep for this. Depth 128-280 m.

Study material.—Paratypes, CARIBBEAN: ANSP 130812 (1: 167 incom-
plete), PILLSBURY 739, 10°54.7'N, 66°17.8’W, 234-280 m, 23 July 1968.
USNM 200779 (1: 151), OREGON 5628, 10°42’N, 67°56’W, 210 m, 28 Sept.
1965.

Paratype, GULF OF MEXICO: USAIC 03460 (1:201), same data as ho-
lotype.

Etymology.—From Gr. pygmaios: small, dwarf, referring to the small
size of the adults. An adjective.

Nettenchelys gephyra, new species, Castle and Smith
Figs. 8B, 11

Holotype.—MNHN 1979-150 (female: 431), Bulari Pass (western reef of
New Caledonia), 400 m, 1 April 1978.

Diagnosis.—Anterior median vomerine teeth enlarged (Fig. 11B). Poste-
rior nostril on top of head about midway between posterior margin of eye
and occiput (Figs. 8B, 11D). Mature at 430 mm TL.

Description (from holotype).—Meristic characters: Total vertebrae 158
(incomplete). Brachiostegal rays 5 on right side. Pores: LL 44; POM 17; IO
oF 82 SOD SST 3:

Morphometric characters: Preanal ca. 39% TL; predorsal 31% preanal;
head 29% preanal; snout 36% head; eye 12% head; upper jaw 51% head;
gill opening 6% head; depth 9% preanal.

Dentition (Fig. 11B, C): Maxillary teeth in bands, those of innermost
series distinctly enlarged; mandibular teeth in bands, those of innermost
series markedly enlarged; premaxillary tooth patch short, continuous with
maxillary teeth; vomerine tooth patch elongate, reaching to level of anterior
margin of eye, anteriormost part isolated from remainder and consisting of
3 enlarged teeth in a single row.

554 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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Fig. 11. Nettenchelys gephyra, holotype (431 mm). A, Head. B, Upper dentition. C, Lower
dentition. D, Dorsal view of a portion of head showing posterior nostrils. Scales in mm; a
single scale applies to B and C.

Color: In preservative light brown, darker on head; posterior portions of
dorsal and anal fins black.

Size: The holotype, a female of 431 mm TL, 167 mm preanal, had eggs
of approx. 0.9 mm diameter. The species probably does not grow much
larger.

Relationships.—This species most closely resembles the Atlantic N. pyg-
maeus. The principal differences are the size at maturity and the position
of the posterior nostril. Whereas the largest N. pygmaeus was 201 mm TL,
and another specimen of 151 mm also contained large eggs, the holotype of
N. gephyra measured 431 mm. The posterior nostril is located more pos-
teriorly in the holotype of N. gephyra than in N. pygmaeus. Nettenchelys
gephyra has slightly more preanal lateral-line pores, one more infraorbital
pore, and a slightly lesser upper jaw length than N. pygmaeus, but these
are based on so few specimens that their significance is uncertain.

The second specimen, USNM 134972, is much smaller, but it is badly
damaged and neither the sex nor the state of maturity can be determined.
Its dentition is identical to that of the holotype, and its posterior nostril is

VOLUME 94, NUMBER 2 555

about a third of the way between the posterior margin of the eye and the
occiput. We assume it is N. gephyra, although because of its poor condition
we do not designate it as a paratype.

Range.—The two specimens were collected off New Caledonia and the
Philippines. Depth about 400 m.

Study material.—USNM 134972 (1: 93 incomplete), ALBATROSS
D5187, 9°16’45’N, 123°21'1S”E, 412 m, 31 March 1908.

Etymology.—Gr. gephyra, a bridge, in reference to the link that this
species provides between Nettenchelys pygmaeus and Nettenchelys inion,
which has the posterior nostrils farther back. A noun in apposition.

Nettenchelys inion, new species, Smith and Bohlke
Figs. 8C, 9

Holotype. —ANSP. 142274 (male: 423), GERDA 242, 25°36'’N, 79°21'W,
458-531 m, 30 Jan. 1964.

Diagnosis.—Posterior nostril at occiput, just in front of supratemporal
canal (Fig. 8C). Median supratemporal pore present. Anterior vomerine
teeth not enlarged. Predorsal 37% preanal, head 38% preanal.

Description (from holotype).—Meristic characters: Vertebrae 210 (com-
plete?). Brachiostegal rays 6 on right side. Pores: LL 41; POM 14; IO 9 +
S12 ONS aus) base

Morphometric characters: Predorsal 37% preanal; head 38% preanal;
snout 37% head; eye 10% head; upper jaw 55% head; depth 10% preanal.

Dentition: Maxillary teeth in bands, innermost teeth somewhat enlarged.
Mandibular teeth in bands, inner teeth enlarged. Premaxillary tooth patch
short, continuous with maxillary teeth. Vomerine tooth patch elongate,
reaching about to eye, anterior teeth not enlarged.

Color: In preservative brown; vertical fins dark-edged posteriorly.

Size: The only known specimen is 423 mm TL, 127 mm preanal.

Relationships.—The single known specimen most closely resembles N.
exorius n. sp. The principal difference is the position of the posterior nostril.
In N. inion the supratemporal canal is unobstructed, and the median pore
is present. The holotype of N. inion has slightly greater predorsal, head,
and snout lengths than any of the four specimens of N. exorius. Since the
specimens of N. exorius include both males and females, these differences
cannot be sexual. There is no evidence elsewhere among the Nettastoma-
tidae that the position of the posterior nostril varies greatly within a species,
and we conclude that the specimen described here represents a distinct
species.

Range.—The single known specimen was collected just south of Bimini
at the eastern edge of the Straits of Florida, depth 458-531 m (Fig. 9).

Etymology.—Gr. inion, back of head, occiput, referring to the position
of the posterior nostril. A noun in apposition.

556 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

OS
——< SRRAdsS

__

S CS OS, S Be
Rete ie Mensa =

; rie lhe CEE SOL

ws 2 3
ARIAS Se a ae AEG”, tee WAMAat,

Fig. 12. Nettenchelys exorius, ANSP 130813 (429 mm). A, Head. B, alone dentition. C,
Lower dentition. Scales in mm; a single scale applies to B and C.

Nettenchelys exorius, new species, Bohlke and Smith
Figs. 8D, 9, 12

Holotype.—ANSP 142275 (female: 467), SILVER BAY 3685, 29°41’N,
80°11’W, 347 m, 22 Jan. 1962.

Diagnosis.—Posterior nostril behind occiput, between supratemporal ca-
nal and dorsal-fin origin (Fig. 8D). Median supratemporal pore absent, ap-
parently obliterated by nasal tube. Anterior vomerine teeth not enlarged
(Fig. 12B). Predorsal 32-34% preanal, head 31-34% preanal.

Description.—Meristic characters: Number of vertebrae unknown; all
specimens have regenerated tails. Branchiostegal rays 6 on right side in one

VOLUME 94, NUMBER 2 S5)//

paratype (ANSP 130813). Pores: LL 40-43 (4); POM 14-16 (3); IO 8-10 +
3 (4); SO 5 (4); ST 2 (4)

Morphometric characters: Predorsal 32-34% preanal (4); head 31-34%
preanal (4); snout 34-36% head (4); eye 9-11% head (4); upper jaw 45-51%
head (3); gill opening 5—7% head (3); interbranchial 15—19% head (2); depth
8—12% preanal (4).

Dentition (Fig. 12B, C): As in N. inion.

Osteology: Supraoccipital absent. Upper pharyngeals divided. First ba-
sibranchial ossified, closely articulated with glossohyal. Second basibran-
chial cartilaginous. Very small epicentral processes on caudal vertebrae.

Color: In preservative, brown; vertical fins dark-edged posteriorly.

Size: The largest specimen was 467 mm TL, 163 mm preanal.

Range.—Three of the four known specimens were collected off the north-
east coast of Florida. The fourth came from the junction of the Northwest
and Northeast Providence Channels in the Bahamas. Depth 277-494 m.

Study material.—Paratypes: ANSP 130813 (1: 429), OREGON 5745,
29°40’N, 80°03’W, 458 m, 18 Nov. 1965. ANSP 131739 (1: 329), OREGON
11703, 30°28'N, 79°51'N, 494 m, 19 Jan. 1972.

Other: UMML 20867 (1:253—cleared and stained), GERDA 867, 25°52'N,
77°54'W, 277 m, 20 July 1965.

Etymology.—Gr. exorios, beyond the frontier, in reference to the pos-
terior nostrils, which have migrated beyond the occiput and are located
behind the head. An adjective.

Nettenchelys taylori Alcock, 1898
Fig. 8E

Nettenchelys taylori Alcock, 1898:150 (original description, off the Trav-
ancore coast, India, holotype listed below); 1899, pl. 25 (Fig. 5).

Diagnosis.—Posterior nostril far behind occiput, posterior to midpoint
between snout tip and anus (Fig. 8E). Median supratemporal pore absent.
Anterior vomerine teeth not enlarged. Predorsal 34% preanal, head 33%
preanal.

Description.—Meristic characters: Pores: LL 48; POM 15; IO 9 + 3/
8 + 3;SO 5; ST 2. Vertebrae: Tail tip regenerated, 48 vertebrae before anal
fin.

Morphometric characters: Predorsal 34% preanal; head 33% preanal;
snout 33% head; eye 8% head; upper jaw 47% head; gill opening 12% head;
interbranchial 15% head; depth at gill opening 8% preanal; depth at anus
10% preanal; snout to posterior nostril 71% of preanal on left side; 68% on
right.

Dentition: Maxillary tooth patches irregularly six teeth across at their

558 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

widest, the teeth of the innermost row moderately enlarged anteriorly but
not posteriorly; teeth gradually increasing in size from lateral to medial.

Premaxillary tooth patch irregularly five or six teeth deep (from front to
rear) at its broadest, the teeth gradually increasing in size from front to rear;
those of posteriormost series enlarged like the anteriormost inner maxillary
teeth and, like them, with arrow-shaped tips. Premaxillary and maxillary
patches continuous without a space between them.

Vomerine tooth patch twelve rows across at its widest, none of the teeth
enlarged, patch terminating posteriorly far short of maxillary and mandib-
ular patches. Teeth largest medially, decreasing in size in both directions
laterally. A rather wide separation anteriorly between premaxillary and vo-
merine patches.

Mandibular tooth patches irregularly about nine teeth across at widest,
the teeth increasing in size from outside inward, those of the innermost
series moderately enlarged except posteriorly; patches distinctly separated
from each other anteriorly. Mandibular patches continue posteriorly to ric-
tus, while maxillary patches extend back beyond that point laterally.

Color: In preservative, brown; vertical fins pale, dusky marginally.

Size: The only known specimen, the holotype, is 533 mm TL. It is a
female with eggs.

Range.—The only specimen was collected at a depth of 786 m off the
Travancore coast, a region in the State of Kerala at the southwestern tip of
India.

Study material.—Holotype: ZSI F 3171/1 (female: 533 incomplete), IN-
VESTIGATOR Sta. 232, 7°17'30’N, 76°54'30"E, 786 m, 19 Oct. 1897.

Etymology.— ‘Named after Commander A. Dundas Taylor, late Indian
Navy, who, in the year 1874 was chiefly instrumental in reviving the Marine
Survey of India’ (Alcock, 1898:151).

Acknowledgments

We thank the following for access to specimens or for information on
them: E. Bertelsen, Zoological Museum, Copenhagen; A. Crosnier,
O.R.S.T.O.M., Paris; P. Fourmanoir, O.R.S.T.O.M., Noumea; J. Paxton,
Australian Museum; D. Baron, Tasmanian Fisheries Development Author-
ity; J. E. Randall, Bernice P. Bishop Museum; P. K. Talwar, Zoological
Survey of India; C. R. Robins, University of Miami; R. L. Shipp, University
of South Alabama; and the curators of the Division of Fishes, U. S. National
Museum of Natural History. We thank R. L. Shipp in addition for permitting
the holotype of Nettenchelys pygmaeus to be deposited at the Academy of
Natural Sciences of Philadelphia. Figures 3, 7 and 11 were done by P. H.
J. Castle; Figures 6, 10B, 10C, 12B and 12C by Mary H. Fuges; and the
remainder by David G. Smith. We thank Eugenia B. Bohlke for X-rays,

VOLUME 94, NUMBER 2 559

stained preparations, and assistance with the literature. C. Karrer informed
us of the presence of Nettastoma solitarium in the western Indian Ocean.
We acknowledge National Science Foundation aid in this study: DEB 76-
20325 (J.E.B.) and INT 77-12568 (D.G.S.). Robert H. Kanazawa indepen-
dently recognized as new the species here named Nettastoma syntresis; we
thank him for relinquishing his material to be included in our revision. This
is a contribution from the Marine Biomedical Institute and the Department
of Human Biological Chemistry and Genetics, University of Texas Medical
Branch at Galveston.

Literature Cited

Alcock, A. 1898. XXII.—Natural history notes from H. M. Indian Marine Survey Ship ‘In-
vestigator,’ Commander T. H. Heming, R. N., commanding.—Series II, No. 25. A note
on the deep-sea fishes, with descriptions of some new genera and species, including
another probably viviparous ophidioid.—Ann. Mag. nat. Hist., (7):136—156.

. 1899. Illustrations of the zoology of the Royal Indian Marine Survey Ship Investigator,
under the command of Commander T. H. Heming, R. N./Fishes.—Part VI, Plates XX V—
XXVI. Crustacea.—Part VII, Plates XXX VI-XLV.—Calcutta.

Costa, O. G. 1844. Fauna de Regno di Napoli. . . Pesci, Pt. 1, Fasc. 48-49, Genere Murena;
Muraena Thunb. (non Artedi): 1-10.

Facciola, L. 1893. Sull’esistenza di forme di passaggio da alcune specie di leptocefalidi agli

adulti corrispondente.—Naturalista Sicil., 12:144—-148.

. 1899. Sull’esistenza di due specie di Nettastoma nel Mare di Messina.—Riv. Ital. Sci.

Nat., 19(2—3):29-32.

. 1911. Generi di vertebrati ittioidi del mare di Messina.—Boll. Soc. Zool. Ital.,

(2)12:268-287.

. 1914. Su di un nuovo tipo dei Nettastomatidi.—Boll. Soc. Zool. Ital., (3)3:39—47.

Gilbert, C. H. 1905. II. The deep-sea fishes of the Hawaiian Islands. In: Jordan, D. S., and
B. W. Evermann, The aquatic resources of the Hawaiian Islands.—Bull. U.S. Fish.
Commn., 23 (for 1903), Part 2:575-713, pls. 66-101.

Gunther, A. 1870. Catalogue of the fishes in the British Museum, 8:xxv + 549 p.

1877. Preliminary notes on new fishes collected in Japan during the expedition of

H.M.S. ‘Challenger.—Ann. Mag. nat. Hist., (4)20:433-446.

Jordan, D. S. 1917-1920. The genera of fishes . . . in four parts.—Stanford University, Uni-
versity Series: 576 pp. (+ total of 55 pp. variously paged in Roman numerals). [Part I,
1917, pp. 1-161; Part II, 1919, pp. 163-284; Part III, 1919, pp. 285-410; Part IV, 1920,
pp. 411-576].

Kaup, J. J. 1856. Catalogue of apodal fish, in the collection of the British Museum.—London:
British Museum, viii + 163 p., 19 pls.

K6lliker, A. 1854. Weitere Bemerkungen tiber die Helmichthyiden.—Verh. Phys.-med. Ges.
Wurzb., 4:100-102 (A. KOlliker in this paper although given as Rudolph Albert von
Kolliker in Dean’s Bibliography of Fishes).

Kotthaus, A. 1968. Fische des Indischen Ozeans. Ergebnisse der ichthyologischen unter-
suchungen... ‘“‘Meteor’’. . . A. Systematischer Teil, III, Ostariophysi und Apodes.—
Meteor Forsch. (D) 3:14—S6.

Lea, E. 1913. Muraenoid larvae from the ‘*Michael Sars’’ North Atlantic deepsea expedition
1910.—Rep. sci. Res. “‘Michael Sars’’ N. Atl. Deep Sea Exped. 1910, 3 (1), Zoology: 1-
59, 6 pls.

560 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Osorio, B. 1909. Contribui¢gao para o conhecimento da fauna bathypelagica visinha das costas
de Portugal/Prefacio.—Mems. Mus. Bocage, 1:1-35, 3 pls.

Poll, M. 1953. Poissons: Téléostéens malacoptérygiens.—Res. sci. Exped. océanogr. belg.
eaux cot. afr. Atl. Sud., 4(2)3:258 p., 104 figs., 8 pl.

Rafinesque, C. S. 1810. Caratteri di alcuni nuovi generi e nuove specie di animali e piante
della Sicilia, con varie Osservazioni sopra i medisimi.—Palermo: 105 p., 20 pls.

Risso, A. 1810. Ichthyologie de Nice, ou histoire naturelle des poissons du département des

Alpes maritimes.—Paris, Schoell: xxxvi + 388 p.

. 1826. Histoire naturelle des principales productions de 1’ Europe méridionale et par-

ticulierement de celles des environs de Nice et des Alpes maritimes. Vol. 3.—Paris and

Strasbourg: xvi + 480 p., 16 pls.

Seeley, H. G. 1874. On Muraenosaurus Leedsii, a plesiosaurian from the Oxford clay.—
Quart. J. Geol. Soc. Lond., 13(21):197—208, pl. 21.

Smith, D. G. 1979. A guide to the leptocephali (Elopiformes, Anguilliformes, and Notacan-

thiformes).—NOAA tech. Rep. nat. mar. Fish. Serv. Fish. Circ. 424: iv + 39 pp.

. and R. H. Kanazawa. 1977. Eight new species and a new genus of congrid eels from

the western North Atlantic with redescriptions of Ariosoma analis, Hildebrandia guppyi
and Rhechias vicinalis.—Bull. mar. Sci. 27(3):530-543.

Whitley, G. P. 1951. New fish names and records.—Proc. R. Zool. Soc. N.S.W., 1949-50:61-—
68.

(DGS) Marine Biomedical Institute, University of Texas Medical Branch,
200 University Blvd., Galveston, Texas 77550; (JEB) Academy of Natural
Sciences, 19th and the Parkway, Philadelphia, Pennsylvania 19103; (PHJC)
Department of Zoology, Victoria University, Private Bag, Wellington, New
Zealand.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 561-563

A NEW GONIASTERID SEASTAR, EVOPLOSOMA
SCORPIO (ECHINODERMATA: ASTEROIDEA),
FROM THE NORTHEASTERN ATLANTIC

Maureen E. Downey

Abstract.—Evoplosoma scorpio, a new species of starfish (Family Go-
niasteridae) from the western approaches to the English Channel, is de-
scribed. It is chiefly characterized by its complete covering of irregular
granules and acute, conical spines, and by tall, spatulate, denticulate pedi-
cellariae.

A new goniasterid starfish collected by the Sarsia in the western ap-
proaches to the English Channel (48°N, 10°W) in ca. 1600 meters represents
a genus hitherto unknown from the Atlantic. Of the two species previously
described in this genus, Evoplosoma forcipifera Fisher (1906) is known from
Hawaii, and E. augusti (Koehler (1909) from the Indian Ocean.

The genus Evoplosoma, described by Fisher in 1906, belongs to the
subfamily Hippasteriinae, of the family Goniasteridae. The other Atlantic
genus in this subfamily, Hippasteria, is abundant north of ca. 40° in both
the eastern and western Atlantic.

Evoplosoma scorpio, new species
Fig. |

Holotype.—Deposited in the British Museum (Natural History).

Type locality.—Sarsia Station 2/19, 48°28.6’N, 10°20.1’W, in western ap-
proaches, west of English Channel and south of Ireland; 1600 m.

Description.—Disc pentagonal, inflated; arms 5, long, narrow, cylindrical;
abactinal plates small, irregularly circular, flat, of varying sizes, completely
covered by irregular, tumid granules (usually with 1 or more tiny conical
‘‘pimples’’) and bearing each 1-2 short, acute, conical spines, often accom-
panied by a small feliped pedicellaria; granules covered with thin skin; abac-
tinal disc surface larger than actinal surface, marginals not visible from
above; marginal plates corresponding throughout length of arm, moderate,
broader than long, rectangular, covered with granules and bearing 1—5 con-
ical spines slightly longer than those on abactinal plates; actinal areas rather
small, with ca. 6 rows of actinal plates, row adjacent to adambulacrals
extending more than halfway out arm; actinal plates irregularly polygonal,
armed as abactinals but with larger, more abudant pedicellariae; adambu-
lacral plates more or less square, furrow margin straight or slightly curved,

562 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Evoplosoma scorpio, abactinal, actinal, and lateral views.

bearing 5—6 stout, blunt furrow spines, somewhat flattened laterally; behind
furrow spines, | large conical subambulacral spine, 1 pedicellaria, several
granules; mouth plates large, rhomboid, bearing ca. 5 large, sturdy, flattened
oral spines on each half, first spine largest, and ca. 2 rows of angular gran-
ules on either side of furrow; pedicellariae taller than broad, spatulate, fe-
lipedal, bearing blunt, spaced teeth across apex and down each side of
valves; madreporite ordinary, of moderate size, closer to margin of disc
than center. R = 110 mm, r = 33 mm, height of disc = 20 mm, number of
superomarginals = 33.

VOLUME.94, NUMBER 2 563

Color.—Bright orange-red, center of disc paler, madreporite white.

Etymology.—Named for the constellation Scorpio.

Discussion.—The form of this species is exactly that of Evoplosoma for-
cipiferum Fisher, from Hawaii, with an abrupt demarcation between the
pentagonal disc and the narrow arms. It differs from that species in having
a much thinner, less conspicuous membrane, in having fairly uniform abac-
tinal spines (not ‘‘spines and spinelets,’’ as Fisher describes E. forcipifera),
and the adambulacral spines are straight, slightly flattened, of the ordinary
goniasterid type, not ‘‘remarkably thin and compressed . . . with expanded
chisel-like tips’’ as in E. forcipifera. The marginal plates, though not large,
are quite distinct, the opposite being true for E. forcipifera. There are twice
as many superomarginal plates in E. scorpio, but this may be due to the
difference in size. The valves of the pedicellariae, somewhat fluted and with
irregular denticulation in Fisher’s species, are much smoother and more
regular in E. scorpio. The actinal interradial areas are smaller, but the first
row of actinal plates extends further out on the arm in E. scorpio than in
E. forcipifera.

Evoplosoma augusti Koehler, from the Indian Ocean, differs in form from
the other two species, being of the more conventional stellate shape, without
the inflated disc; Koehler says (1909:96): ‘‘edge of disc notably thinner than
center.’ The abactinal plates bear fat tubercles, rather than conical spines,
and there is a bare space between the tubercles and the surrounding gran-
ules. As in E. forcipifera, the actinal areas are larger than in E. scorpio.
Evoplosoma augusti has ca. 8 truncate, lamelliform adambulacral spines;
the pedicellariae are exactly like those of E. forcipifera. The peculiar body
form of Evoplosoma scorpio is also not uncommon in north Atlantic spec-
imens of Hippasteria phrygiana, to which the genus Evoplosoma is closely
related.

Acknowledgments

Thanks are due to Ailsa M. Clark, of the British Museum (National His-
tory), and Eve Southward, of the Plymouth Laboratory, for allowing me to
examine and describe this species. I thank Miss Clark also for the photo-
graphs, and for reading the manuscript.

Literature Cited

Fisher, W. K. 1906. The starfishes of the Hawaiian Islands.—Bull. U.S. Fish Comm. 1903,
23(3):987—-1130, pls. 1-49.

Koehler, R. 1909. Astéries recueillies par Investigator dan l’Ocean Indien. I. Les Astéries
de Mer profonde.—Calcutta: Indian Museum, Pp. 5-143, pls. I-XIII.

Department of Invertebrate Zoology, National Museum of Natural His-
tory, Smithsonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 564-568

A REDESCRIPTION OF TUBIFICOIDES
HETEROCHAETUS (MICHAELSEN)
(OLIGOCHAETA: TUBIFICIDAE)

H. R. Baker

Abstract.—Tubificoides heterochaetus (Michaelsen, 1926) (Oligochaeta:
Tubificidae) is redescribed from type-material and is compared to new ma-
terial of this species from North Carolina. Oversights in the original de-
scription and recent literature are corrected.

Access to one of the syntypes of Tubificoides heterochaetus (originally
described as Limnodrilus heterochaetus Michaelsen, 1926), and to mature
material of this species from North Carolina has necessitated a redescription
as the original description and recent literature have been found to be lack-
ing in some respects.

This species is now seen to possess a thin but distinct penis sheath and
both bifid and simple-pointed setae posteriorly.

The syntype described here is accorded lectotype status.

Methods and Materials

One of the syntypes of Limnodrilus heterochaetus Michaelsen, 1926 was
loaned to Dr. R. O. Brinkhurst, courtesy of Dr. M. Dzwillo, Zoologisches
Institut and Zoologisches Museum (ZIZM), Universitat Hamburg, W. Ger-
many. Mr. Brian Woodard sent several specimens of 7. heterochaetus from
North Carolina for identification. All material was stained in paracarmine
and mounted whole in Canada Balsam. Some of the North Carolina material
has been deposited in the Smithsonian Institution, United States National
Museum of Natural History (USNM), Washington, D.C.

Systematic Description

Tubificoides heterochaetus (Michaelsen, 1926)
Figs. 1, 2

Limnodrilus heterochaetus Michaelsen, 1926:22, fig. A (a—d); 1927:17,
fig. 19.

Limnodrilus heterochaetus Michaelsen.—Btilow, 1957:99.—Popescu-Mari-
nescu et al., 1966: 1963.

Peloscolex heterochaetus (Michaelsen).—Marcus, 1942:156.—Marcus,
1950:4.—Cekanovskaya, 1962:279, figs. 176, 177.—Brinkhurst, 1963a:44,

VOLUME 94, NUMBER 2 565

C 15um

A 100 um

Fig. 1. Tubificoides heterochaetus: A, Male duct and spermatheca of lectotype, notation
as in description, penis sheath thickened slightly to show form; B, Dorsal anterior bifid seta;
C, Dorsal posterior bifid seta.

fig. 31; 1963b:713.—Hrabé, 1964:109.—Brinkhurst and Jamieson, 1971:521,
figs. 8, 14 (G_I).
Tubificoides heterochaetus (Michaelsen).—Brinkhurst and Baker, 1979:1556.

Description (lectotype).—Length 5.1 mm, width at X 380 um (in fixed,
slightly compressed specimen), 29 segments (incomplete). Prostomium
shorter than broad at peristomium, conical. Clitellum weak over 2X—Y“XII.
Body wall slightly papillate posteriorly with foreign material adhering. Seg-
ments II-VIII short, segment IX longer, gut enlarged in IX.

Anteriorly 4, 5 bifid setae per bundle, upper tooth about as long and thick
as lower (Fig. 1B); 1, 2 setae per bundle posteriorly, one ventral bundle of
X with | seta (other side missing seta), no ventral setae in XI. No modified
genital setae. From XIV—XV posteriorly ventral setae sharply simple-point-
ed; some (all?) posterior dorsal bundles with bifid and simple-pointed setae,
bifid setae with upper tooth longer and much thicker than the very thin lower
tooth (Fig. 1C). Male and spermathecal pores in setal line, spermathecal
pores open anterior to ventral setae of X.

Male genitalia (all structures paired) (Fig. 1A): sperm funnel (sf) moderate
in size, vas deferens (vd) 35.5 wm wide behind sperm funnel narrowing to
16.9 um just before entry into atrium (at); vas deferens about as long as

566 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

atrium with wide ciliated lumen. Atrium with enlarged muscular (m) head,
vas deferens and prostrate (pr) enter nearly opposite each other just below
swollen head; prostate small, stalked. Atrium apparently tripartite with
swollen head being composed of large (vacuolated?) cells, middle tubular
portion composed of large granular cells, lower (distal) portion composed
of small granular cells. Atrium narrows basally and then widens just before
joining penial bulb. Atrium terminates in true penis (p) with thin cuticular
sheath; penis sheath (ps) short (46 wm long) and wide (approx. 36 wm wide
at base), slightly cone shaped, in short penial sac (psc).

Spermathecae (s) short (Fig. 1A) (approx. 200 um in length) with thick
walled ampullae and distinct ducts (approx. 35 wm wide), sperm traps (st)
apparently present; basal end of duct bulbous (46 um in diameter) with
distinct lumen. No spermatozeugmata observed.

Material examined.—ZIZM cat. V9920. Lectotype of T. heterochaetus,
whole mounted specimen.—USNM 62805, 62806, 2 whole mounted specimens
from Cape Fear River (32 miles from coast), North Carolina. Sample from
25 ft, locality estuarine, medium sands with silt and organic detritus. B.
Woodard coll., 28 Feb. 1980. Found with Dero sp., Paranais sp., Nais
communis, Nais variabilis (all naidid oligochaetes), Scolecolepides viridis
(a spionid polychaete) (B. Woodard, pers. comm.).—Baker collection, 5
whole mounted specimens, locality and collector as for USNM speci-
mens.—Woodard collection, 3 whole mounted specimens, locality and col-
lector as for USNM specimens.

Distribution.—Europe, North Carolina, Virginia, U.S.A.

Discussion

The description of Tubificoides heterochaetus given here differs from the
original (Michaelsen, 1926) in one important respect. In the original descrip-
tion the dorsal posterior setae were described as being sharply simple pointed.
The lectotype of T. heterochaetus has some posterior dorsal setae which are
clearly bifid (Fig. 1C). The lower tooth of these setae is very thin, rather
short, and is not visible if the setae are not properly oriented.

Some of the recent literature (Brinkhurst and Jamieson, 1971; Brinkhurst
and Baker, 1979) reported T. heterochaetus to be without a penis sheath.
Although Michaelsen (1926:27, 28) did recognize that 7. heterochaetus had
a slightly thickened cuticular penis sheath he felt that it was not thick or
stiff enough to be called a true penis sheath. T. heterochaetus is recognized
here as possessing a true penis sheath.

The specimens of 7. heterochaetus from North Carolina agree for the
most part with the above description. These specimens reach a maximum
length of 9 mm with up to 31 segments. There may be up to 3 setae per
bundle posteriorly. Sperm traps (sensu Brinkhurst and Baker, 1979) are

VOLUME 94, NUMBER 2 567

100 um

| Fig. 2. Tubificoides heterochaetus: Spermathecae of a North Carolina specimen, notation as
in description, sperm trap clearly present, basal portion of spermathecal duct bulbous with
distinct lumen.

present in the North Carolina specimens (Fig. 2). As the lectotype had not
yet mated (shown by lack of Spermatozeugmata in the spermathecae) this
feature was not as apparent. The spermatozeugmata (sp) in the North Car-
Olina heterochaetus are elongate in shape (Fig. 2). One or both of the Sper-
mathecal ampullae often penetrate the sperm sac into XI. In most of the
North Carolina heterochaetus there is 1 seta present per ventral bundle
of X.

In the North Carolina specimens the atrial muscle layer below the head
of the atria does not thin out as fast and remains slightly thicker than does
the muscle layer in the lectotype (Fig. 1A). This may be due to different
fixing procedures in the material from the two locations.

The North Carolina heterochaetus also have bifid dorsal setae posteriorly
which are of the same form as those of the lectotype. This is a very unusual
characteristic in the Tubificinae as the upper tooth is usually the tooth
which undergoes reduction. The presence of this feature in both the lecto-
type and the North Carolina heterochaetus is a very strong indication that
the North Carolina material is indeed T. heterochaetus. Many of these dor-
Sal posterior bundles consist of 1 bifid seta with the other setae being
sharply simple-pointed setae.

_ The North Carolina heterochaetus are similar to the lectotype in their
degree of papillation.

|
|
|

568 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

All members of the genus Tubificoides (sensu Brinkhurst and Baker,
1979) are now known to possess a penis sheath of some form.

Acknowledgments

I thank Dr. M. Dzwillo and Mr. B. Woodard for making available the
material used in this study. Dr. R.O. Brinkhurst’s encouragement and crit-
icism of this manuscript is appreciated. This work was supported by a Brit-
ish Columbia Provincial Government G.R.E.A.T. award (with the cooper-
ation of Dobrocky Seatech Ltd.).

Literature Cited

Brinkhurst, R. O. 1963a. Taxonomical studies on the Tubificidae (Annelida, Oligochaeta).—

Int. Revue Ges. Hydrobiol. 48:7-89.

. 1963b. Notes on the brackish-water and marine species of Tubificidae (Annelida,

Oligochaeta).—J. Mar. Biol. Ass. U.K. 43:709-715.

, and H. R. Baker. 1979. A review of the marine Tubificidae (Oligochaeta) of

North America.—Canadian J. Zool. 57:1553-1569.

, and B. G. M. Jamieson. 1971. Aquatic Oligochaeta of the World.—Oliver and

Boyd, Edinburgh. 860 pp.

Bulow, T. 1955. Oligochaeten aus den Endgebieten der Schlei.—Kieler Meeresforsch. 11:253-
263.

Cekanovskaya, O. V. 1962. The aquatic Oligochaete fauna of the USSR.—Opred. Faune
SSSR 78:1-411. [in Russian]

Hrabe, S. 1964. On Peloscolex svirenkoi (Jaroschenko) and some other species of the genus
Peloscolex.—Spisy Prir. Fac. Univ. Brne 450:101-112.

Marcus, Ernesto. 1942. Sobre algumas Tubificidae do Brasil.—Bolm. Fac. Filos. Cienc. Univ. |
Sao Paulo 25:153-228.

Marcus, Eveline du B.-R. 1950. A marine tubificid from Brazil. Commun. Zool. Mus. Hist.
Nat. Montevideo 3(59):1-8.

Michaelsen, W. 1926. Oligochaeten aus dem Ryck bei Greifswald und von benachbarten

Meeresgebieten.—Mitt. Hamb. Zool. Mus. Inst. 42:21-29.

. 1927. Clitellata: Oligochaeta.—In: Grimpe and Wagler, Die Tierwelt der Nord- und

Ostsee (chap. VI c:1-44), Leipzig.

Popescu-Marinescu, V., F. Botae, and G. Brezeanu. 1966. Untersuchungen tiber die Oligo- |
chaeten im rumanischen Sektor des Donaubassins.—Arch. Hydrobiol. Suppl. 30:161-
179.

Department of Biology, University of Victoria, Victoria, B.C., Canada |
V8W 2Y2.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 569-578

THE GENERIC STATUS AND DISTRIBUTION OF
MONODELLA TEXANA MAGUIRE, THE ONLY
KNOWN NORTH AMERICAN THERMOSBAENACEAN

Jan H. Stock and Glenn Longley

Abstract.—Although the original description of Monodellu texana Ma-
guire, 1965, attributes several peculiar features, notably in the maxillipeds,
to the sole North American representative of the Thermosbaenacea, a re-
examination of material from the San Marcos area (Texas) revealed a close
morphological similarity to European and West Indian taxa of Monodella,
with which it clearly is congeneric.

Maguire’s (1964, 1965) discovery of the first thermosbaenacean outside
the Mediterranean area awoke considerable interest by biogeographers and
stygobiologists. Maguire attributed his material without comments to the
genus Monodella, at that time only known from groundwaters in Italy,
Yugoslavia, and Israel. Presently, the genus is known also from Spain, the
Balearic Islands, France, and Greece, and outside the Mediterranean area
from the British and U.S. Virgin Islands, Culebra (E. of Puerto Rico), Haiti,
Cuba (references in Stock, 1976, and Stock, in press), Puerto Rico (unpub-
lished personal observations) and Somalia (Messana, 1979).

In the years following the description of the Texan thermosbaenacean,
but preceding the discovery of extra-Mediterranean taxa, several carcinol-
ogists wondered whether Maguire had been right in considering the New
World taxon congeneric with the Old World species. When Stock (1976)
described a second New World species, found in St. Croix (U.S. Virgin
Islands), it became clear that true Monodellas existed outside the Medi-
terranean, but at the same time doubt was cast on the correctness of Ma-
guire’s morphological observations. According to the original description,
the Texan Monodella had a 2-segmented mandibular palp (versus 3-seg-
mented in the other species), was devoid of endo- and exopodites on the
2nd maxilla (versus present), and fused coxo- and basipodal endites in the
maxilliped (versus separate), and lacked maxillipedal epipodites (versus pres-
ent). The most conspicuous difference between Maguire’s description of M.
texana and the other members of Monodella was the alleged presence, in
the female, of a 2-segmented maxillipedal endopodite, whereas the normal
female condition is characterized by the absence (or reduction to a vestigial
setule) of an endopodite. The male maxillipedal exopodite of M. texana was
described as 4-segmented, whereas 2-segmented is the normal situation.

570 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

None of these differences materialized in a new study based on 15 spec-
imens from an artesian well in San Marcos, Hays County, Texas, not far
from the type-locality. Since the original description obviously is wanting,
and since the accompanying illustrations are on much too small a scale to
make recognition of details possible, Monodella texana is completely re-
described in the sequel.

The artesian well from which the present specimens came is at an old
Federal Fish Hatchery, deeded to the Southwest Texas State University as
an Aquatic Station in 1964. It derives its water and very interesting hypo-
gean fauna from the underlying Edwards Aquifer. More details and an il-
lustration of the artesian well can be found in the recent paper on the Am-
phipoda of the well (Holsinger and Longley, 1980).

The specimens have been collected by the junior author and his team at
the Edwards Aquifer Research and Data Center, Southwest Texas State
University, San Marcos, Texas. The senior author is indebted to Dr. John
R. Holsinger, Old Dominion University, Norfolk, Virginia, for bringing the
existence of freshly collected material to his notice.

Monodella texana Maguire, 1965

Monodella: Maguire, 1964:931-—932, fig. 1.

Monodella texana Maguire, 1965:149-154, figs. 1-3, pl. III; Karnei, 1978:38,
fig. 15.

Monadella (lapsus calami) texana: Longley, 1978:23.

Body length 1.6—2.0 mm (<6) or 1.7—2.2 (2). Females differ in external
morphology from males in only a few characters: (1) the maxilliped is devoid
of an endopod; (2) a penis on pereiopod 7 is absent; (3) the proximal fla-
gellum segments of the first antenna are devoid of aesthetes; (4) in certain
phases of life, a dorsal brood pouch is present.

The animal is similar in body shape to other members of the genus Mon-
odella and its appendages are remarkably similar to those of the only other
Eastern Hemisphere species named so far, M. sanctaecrucis Stock, 1976.
In the following description, M. texana will be compared with M. sanctae-
Crucis.

The first antenna (Fig. 1) has a 3-segmented peduncle (protopodite) and
2 flagellae; the main flagellum (exopodite) is 7- to 8-segmented, the acces-
sory flagellum (endopodite) is about 7% the length of the main flagellum and
consists of 5 segments. All segments of the main flagellum in 6 bear very
long, stalked, aesthetes; in @ the proximal flagellum segments are devoid
of aesthetes. The peduncle segments in M. texana are only slightly longer
than wide (versus at least twice as long as wide in M. sanctaecrucis). The
peduncle is armed with several long, plumose setae and some short, naked

VOLUME 94, NUMBER 2 571

Figs. 1-10. Monodella texana: 1, First antenna, 3 (scale AB); 2, Second antenna, d (AB);
3, Left corpus manibulae, 2 (AC); 4, Right mandible, 9° (AC); 5, Paragnath, 2 (AC); 6, First
maxilla, ¢ (AD); 7, Second maxilla, ¢ (AD); 8, Labium, 9 (AC); 9, First pleopod, 3 (AB);
10, Second pleopod, ? (AB). (For scales see Figs. 11-15.) b.e. = basipodal endite, b.e./, b.e.

= first and second basipodal endite; c.e. = coxopodal endite; c.m. = corpus mandibulae;
en = endopodite; ex = exopodite; /.m. = lacinia mobilis; p.i. = pars incisiva:; p.m. = pars
molaris.

SZ PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

setae. The third peduncle segment bears a distal, triangular process armed
with 3 short setules.

The second antenna of M. texana (Fig. 2) is similar to that of M. sanc-
taecrucis. It consists of 5 peduncle segments and 5 flagellum segments.

The mandible consists of a corpus mandibulae and 3-segmented palp (Fig.
4). Palp segment | is squarish and unarmed; segment 2 is elongate and bears
1 subdistal, plumose seta; segment 3 is slightly curved and bears 2 distal
plumose setae, | thin and 1 heavy distal naked setae, and a varying number
(3 to 6) lateral plumose setae. In the corpus mandibulae one can distinguish
a pars incisiva, a lacinia mobilis and a pars molaris; the latter two are
separated by a row of 3-4 plumose and 2 (right mandible) or 3 (left mandible)
naked setae. The lacinia mobilis bears 5 fine teeth (right) or 3 coarse teeth
(left), and the pars incisiva bears 3 pointed teeth (right) or 5 obtuse teeth
(left) (Figs. 3 and 4).

The first maxilla (Fig. 6) shows (1) a coxopodal endite, armed with 3
medial, plumose and 1 medial, naked setae, 6 plumose distal setae, and a
lateral row of cilia; (2) a basipodal endite, distally armed with 6, sawlike
spines, and (3) a ‘‘palp’’ (endopodite) consisting of 3 articles, the second
armed with 2 setae and | large toothed spine, the third with | seta and 1
large toothed spine.

As usual in the genus, the second maxilla (Fig. 7) is complexly built: (1)
a coxopodal endite with 2 medial rows of naked setae and a distal row of
8 plumose setae; (2) a first basipodal endite armed with a distal row of about
16 long, slightly S-shaped spines, distomedially provided with a grasping
edge; (3) a second basipodal endite armed with 7 similar, transformed,
though longer and heavier, spines; and (4) a palp of 2 segments, the basal
one of which carries a rudimentary, bud-like exopodite armed with | seta,
and a unimerous endopodite armed with 4 setae. In M. sanctaecrucis the
endopodite is 2-segmented and carries 7 setae.

The labium (Fig. 8) is a deeply cleft, ciliated lobe.

The maxilliped is sexually dimorphic. In the male, a 5-segmented endop-
odite is present; in the female this is reduced to a single setule (Figs. 11,
12). Furthermore, the appendage shows (1) a small coxopodal endite armed
with 3 plumose setae; (2) a large basipodal endite armed with a row of 4
shorter plumose setae and a row of 8 to 9 longer plumose setae; (3) a
2-segmented exopodite armed with 3 distal plumose setae; and (4) a ciliated
epipodite. In M. sanctaecrucis the exopodite is armed with 3 distal, 1 lateral
and | medial setae.

The first pereiopod (Fig. 13) has a 3-segmented exopodite; in some spec-
imens, the segmentation line between the second and third segments tends
to become indistinct. The second exopodal segment bears 3 medial, plumose
setae; the third segment bears 2 terminal and 2 subterminal plumose setae;

VOLUME 94, NUMBER 2 573

Figs. 11-15. Monodella texana: 11, Maxilliped, 2; 12, Maxilliped, 5; 13, First pereiopod,
2; 14, Uropod, ¢; 15, Telson, ¢ (all to scale AB). b = basis; b.e. = basipodal endite; c =
carpus; c.e. = coxopodal endite; d = dactylus; en = endopodite; ep = epipodite; ex = ex-
opodite, i = ischium; m = merus; pr = propodus; r = rudiment of endopodite; uv = ungulus.

574 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

the lateral armature is reduced to 2 vestigial setules. (In M. sanctaecrucis
the lateral armature consists of 2 plumose setae.) The ischial segment of the
endopodite is almost completely fused with the basipodite (articulated in
M. sanctaecrucis). The shape and armature of the merus, carpus, propodus,
dactylus and ungulus are very similar to that of M. sanctaecrucis.

The second pereiopod (Fig. 16) has a 2-segmented exopodite, lacking any
lateral armature (in M. sanctaecrucis it carries 2 plumose setae). Club-
shaped spines occur on the first exopodite segment (2 spines), the endopodal
propodus (5 spines) and dactylus (5 spines). Fig. 17 shows an aberrant ap-
pendage in which the ungulus and the subterminal dactylar spine are sub-
equal.

The third pereiopod resembles the second; the number of club-shaped
spines on the endopodal propodus and dactylus are 11 and 9, respectively.

The fourth pereiopod is rather similar, too (Fig. 18), but bears 12 club-
shaped spines on the endopodal propodus and 7 such spines on the dactylus.
In M. sanctaecrucis the dactylus bears only 2 spines, and the lateral exo-
podal armature (absent in M. texana) consists of 1 plumose seta.

The fifth pereiopod (Fig. 19) has a relatively short second exopodite seg-
ment, armed with 5 plumose setae (6 in M. sanctaecrucis). The endopodal
propodus bears 16, and the dactylus 8 club-shaped spines (6 and 2 spines,
respectively, in M. sanctaecrucis).

In the sixth pereiopod (Fig. 20), the exopodite is reduced to a single
segment, armed with 4 medial club-shaped spines, 2 medio-subdistal plumose
setae, 1 distal rudimentary seta, and | lateral rudimentary seta. The distal
and subdistal elements are placed on a low projection (‘‘socle’’), which is
lacking in M. sanctaecrucis. The endopodal carpus bears usually 2, some-
times 1, medial setae. The propodus bears 19-20, the dactylus 5-6 club-
shaped spines (in M. sanctaecrucis the dactylus bears only 2 spines).

The seventh pereiopod (Fig. 21) has a thin, one-segmented exopodite,
armed as in P6. The endopodal carpus bears 1 medial seta. The armature
of the propodus and dactylus is as in P6. This appendage differs from that
of M. sanctaecrucis in the presence of socles for the (sub)distal exopodal
elements, and in the higher number of endopodal dactylus spines. In the
male, the coxopodite carries a lateral, curved, finger-shaped appendage, the
penis, which is almost as long as the exopodite (shorter in M. sanctaecru-
ciS).

The first and second pleopods (Figs. 9, 10) are small, finger-shaped, one-
segmented appendages, armed with 5 setae.

The uropod (Fig. 14) has a 2-segmented exopodite and a 1-segmented
endopodite and is very similar in morphology to that of M. sanctaecrucis.

The telson (Fig. 15) bears 2 longer and 1 shorter spine on either side, and
a triangular mid-distal lobe in between the two groups of spines. The anus is

VOLUME 94, NUMBER 2 D5

Figs. 16-19. Monodella texana: 16, Second pereiopod, 2; 17, Endopodal dactylus of aber-

rant second pereiopod, 2; 18, Fourth pereiopod, ?; 19, Fifth pereiopod, 2. (All to scale AB,
see Figs. 11-15.)

576 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

fo

Z.
¢
L

WA
Wy
ee

21 20

Figs. 20-21. Monodella texana: 20, Sixth pereiopod, 2; 21, Seventh pereiopod, ¢ (both
to scale AB, see Figs. 11-15.) pe = penis.

subterminal. In M. sanctaecrucis, the anus is terminal and the distal tri-
angular lobe appears to be lacking.

Remarks.—As the above description shows, M. texana is morphologi-
cally very similar to M. sanctaecrucis. The main differences are (1) the
degree of elongation of the peduncular segments of antenna 1; (2) the num-
ber of setae on the 2nd exopodite segment of the maxilliped; (3) the shorter
endopodite of maxilla 2 in M. texana; (4) the absence of plumose lateral
setae in the exopodite of pereiopods 1 to 5 in M. texana: (5) the shape of
the exopodite of pereiopods 6 and 7; (6) the length of the penis in relation
to the length of the 7th pereiopodal exopodite; (7) the number of spines on
the dactylus of the posterior pereiopods; and (8) the shape of the telson.

The overall resemblance, even in the finer details, to the Old World
species (see, for instance, Rouch, 1965, for good illustrations of Monodella
argentarii Stella, 1951) is likewise striking.

In conclusion, it can be said that the members of the genus Monodella,
notwithstanding their occurrence in several isolated areas of the world, have
retained a remarkable morphological uniformity. So far, no marine repre-
sentatives referable to Monodella are known, although marine Thermos-
baenacea belonging to other genera have recently been found in the West

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VOLUME 94, NUMBER 2

578 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Indies (Stock, 1976, and unpublished). The absence of marine Thermos-
baenacea in the Mediterranean, the only other area in the world where they
have been actively looked for, may be explained by the late Miocene hy-
drographic history of that basin (Stock, 1980; Danielopol, 1980). The sea
level dropped very considerably and much of the remaining water was tem-
porarily transformed into brine. It is conceivable that many marine ances-
tors of the actual stygofaunal elements became extinct in the Mediterranean
during the late Miocene. In the West Indies, where no such drastic salinity
crisis took place, marine Thermosbaenacea could and did, in fact, survive.

Distribution.—The type-locality of M. texana is Ezell’s Cave, in San
Marcos, Hays County, Texas (Maguire, 1964, 1965). Karnei (1978) records
the species also from the Verstraeten Well No. 1 and the City Water Board
Artesia Pump Station Well (both in Bexar Co., Texas). Longley (1978) rec-
ords the species from the artesian well of the Southwest Texas State Uni-
versity Aquatic Station in San Marcos (Hays Co.). Material of the latter
well has been used for the above redescription. Moreover, 7 specimens from
the George Ligocky Farm Well No. H-5-158 (Uvalde Co., Texas) were
studied, but, unfortunately, all these specimens were heavily damaged. As
far as could be ascertained, this material appears to be identical to that of
San Marcos. The known distribution is shown in Fig. 22.

Literature Cited

Danielopol, D. 1980. An essay to assess the age of the freshwater interstitial Ostracoda of
Europe.—Bidr. Dierk., Amsterdam 50(2):V243-291.

Karnei, H. 1978. A survey of subterranean aquatic fauna of Bexar County, Texas.—M.Sc.
Thesis, Southwest Texas State University, San Marcos: 1-118.

Longley, G. 1978. Status of Typhlomolge (=Eurycea) rathbuni, the Texas Blind Salaman-
der.—Endangered Species Rep., U.S. Fish & Wildlife Serv., Albuquerque, New Mexico
2:i—vi, 1—45.

Maguire, B., Jr. 1964. Crustacea: A primitive Mediterranean group also occurs in North

America.—Science 146:93 1-932.

. 1965. Monodella texana n. sp., an extension of the range of the crustacean order

Thermosbaenacea to the Western Hemisphere.—Crustaceana 9(2):149-154, pl. III.

Messana, G. 1979. Groundwater research by the “‘Centro di Studio per la Faunistica ed
Ecologia tropicali’’ in Somalia.—Stygo News 2(2):5-8.

Rouch, R. 1965. Contribution a la connaissance du genre Monodella (Thermosbaenacés).—
Ann. Spéléol. 19(4):717-727.

Stock, H. H. 1976. A new genus and two new species of the crustacean order Thermosbae-

nacea from the West Indies.—Bidr. Dierk., Amsterdam 46(1):47-70.

. 1980. Regression model evolution, as exemplified by the genus Pseudoniphargus

(Amphipoda).—Bijdr. Dierk., Amsterdam 50(1):V105—144.

. The influence of Hadziid Amphipoda on the occurrence and distribution of Thermos-

baenacea and Cyclopoid Copepoda in the West Indies.—Int. J. Speleol. 10(4) [in press].

(JHS) Institute of Taxonomic Zoology, University of Amsterdam, The
Netherlands; (GL) Edwards Aquifer Research and Data Center and Aquatic
Station, Southwest Texas State University, San Marcos, Texas 78666.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 579-590

CAECIDOTEA SALEMENSIS AND C. FUSTIS,
NEW SUBTERRANEAN ASELLIDS FROM
THE SALEM PLATEAU (CRUSTACEA: ISOPODA: ASELLIDAE)

Julian J. Lewis

Abstract.—Two species of Caecidotea belonging to the asellid Hobbsi
Group are described from the Salem Plateau Section of the Ozark Plateau
Province. Caecidotea salemensis is a widespread and common inhabitant
of subterranean habitats in the central part of the Salem Plateau in both
Arkansas and Missouri. Caecidotea fustis is known only from subterranean
habitats in the northeastern part of the plateau.

The Salem Plateau is a section of the Ozark Plateau physiographic prov-
ince, which comprises parts of southern Missouri, northern Arkansas and
western Illinois (Fig. 1). Several thousand caves and springs are known
from the Ordovician limestones and dolomites prevalent in the region
(Bretz, 1956; Bretz and Harris, 1961; Vineyard and Feder, 1974), from which
four subterranean species of Caecidotea are currently known: C. antricola
Creaser (1931), C. dimorpha Mackin and Hubricht (1940), C. extensolin-
guala (Fleming, 1972) and C. serrata (Fleming, 1972). Records of Caecido-
tea stiladactyla Mackin and Hubricht (1940) from the central Missouri part
of the Salem Plateau are probably erroneous determinations of C. fustis.
Two subterranean species are added to the list of the Salem Plateau isopod
fauna herein, an accomplishment made possible largely by the collections
of Leslie Hubricht (LH), James E. Gardner (JEG) and John L. Craig (JLC).

Caecidotea salemensis, new species
Figs. 2-5

Asellus tridentatus.—Fleming, 1972:254 (in part). A. spp.—Craig, 1977:83,
85, 87 (in part).

Material examined.—ARKANSAS: Lawrence Co., Imboden, deep cis-
tern, 29 Aug. 1925, Byron C. Marshall, 61 ¢6¢, 12 22 (USNM 59263).
MISSOURI: Boone Co., stream in Hunter Cave, 5 mi NNW Ashland, 4
Aug. 1940, LH, 4 66, 20 2 2. Carter Co., Norris Cave, small quiet pools,
10 mi NW Van Buren, 29 Jan. 1979, JEG, 2 6 ¢. Dallas Co., Saltpeter Cave,
22 Nov. 1974, O. Hawksley, 4¢6¢,7 22. Douglas Co., seep, on bluff above
North Fork of White River, W. of Roosevelt, 1 May 1940, LH, 19 dd, 22
2 2; small spring on bluff near North Fork of White River, 7 mi E Richville,
1 June 1935, LH, 3 d646,3 29. Franklin Co., drip pools and stream, Mush-

580 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

room Cave, 2.5 mi E. Sullivan, 5 Apr. 1942, LH, 25 66, 11 22; same
locality, 1 June 1974, JLC, 1 6, 1 9. Hickory Co., seep 5 mi W Urbana,
16 Mar. 1975, JLC, 21 juv, 3 2 2; same locality, 23 Mar. 1975, 3 2°. Pettis
Co., Hughesville, cistern, 20 Aug. 1932, A. Gurloff, 6 dd, 5 22 (USNM
67700). Phelps Co., Lane Cave, small quiet pools, 1 mi N Yancy Mills, 17
Jan. 1979, JEG, 11 6d, 8 22; Little Piney Cave, 7 mi SW Rolla, 15 May
1980, JEG, 1 6, 2 2 2; Zorumski Cave, 5 mi WSW Newburg, 17 Mar. 1980,
JEG,9 66,2 2. Pulaski Co., Little Cave, beneath stones in riffles, 1 mi
E. Devils Elbow, 6 Dec. 1978, JEG, 9 dd, 6 22; same locality, 6 Aug.
1979, Julian J. Lewis, Teresa M. Lewis, JEG, 4 ¢d¢, 2 22; York Cave,
drip pools, 1.5 mi N Hanna, 28 Mar. 1942, LH, 5 66, 1 9; same locality,
21 May, 1978, JLC, 1 6, 1 2; McCann Cave #1, 4 mi W Waynesville, 6
Mar. 1976, JLC, 5 646,722. Texas Co., Bat Cave, in small stream, 7 mi
NE Success, 5 July 1940, LH, 1 d, 4 2 2 (USNM 108585); Unnamed Cave
#15, 12 mi W Licking, 21 Apr. 1980, JEG, 9 63,5 92. Washington Co.,
Hamilton Cave, drip pool, 5.5 mi SE Sullivan, 20 July 1940, LH, 6 64,
Do Oe.

A 12.5 mm male from Imboden, Arkansas is the holotype (USNM 59263),
the other specimens from this locality are paratypes (USNM 181300). All
of the material examined has been deposited in the National Museum of
Natural History, Smithsonian Institution.

Description.—Eyeless, unpigmented. Longest male, 16.0 mm, body slen-
der, linear, about 6.9x as long as wide, coxae visible in dorsal view; longest
female, 9.8 mm. Margins of head, pereonites and telson very setose. Head
about 1.8 as wide as long, anterior margin concave, rostrum lacking; post-
mandibular lobes moderately produced. Telson about 1.4 as long as wide,
sides subparallel, caudomedial lobe moderately produced, broadly rounded.

Antenna | reaching middle of last segment of antenna 2 peduncle, flagel-
lum of male of about 17-18 segments, esthete formula 3-0-1-0-1 (Fig. 2d);
flagellum of female shorter, of about 11-12 segments. Antenna 2 reaching
pereonite 7, last segment of peduncle about 1.3 length preceding segment,
flagellum of about 70-84 segments.

Mandibles with 4-cuspate incisors and lacinia mobilis; spine row with
about 14 spines in left, 17 spines in right mandible. Palp with plumose setae
on distal segments. Maxilla 1, apex of outer lobe with 13 robust spines,
inner lobe with 5 apical plumose setae. Maxilliped with about 7 retinacula.

Male pereopod 1 propus about 1.3 as long as wide; palm proximally
with 1—2 small, robust spines, medial process subtriangular, separated from
smaller rounded or slightly bicuspid distal process by U-shaped cleft (Figs.
3a; Saf); dactyl flexor margin with up to 5-6 spines, decreasing to 0—1 in
mature individuals, and rounded process (Fig. 3d). Female pereopod 1 pro-
pus more slender, about 1.4x as long as wide, palm with 2 proximal spines,
processes lacking; dactyl flexor margin with about 5 spines. Pereopod 4
more robust in male than female.

VOLUME 94, NUMBER 2 581

Fig. 1. The Salem Plateau Section (stippled around margins) of the Ozark Plateau Province,
showing the ranges of Caecidotea salemensis (squares) and C. fustis (triangles); stippled areas
within the Salem Plateau indicate areas where non-cavernous rocks occur.

Male pleopod 1 larger than pleopod 2; protopod about 0.7x length of
exopod, with 3-4 retinacula; exopod about 2x as long as wide, with long
plumose setae on distal margin and short setae on concave distal part of
lateral margin. Male pleopod 2 exopod, proximal segment with 0-4 setae,
distal segment suboval, bearing about 15 plumose setae along lateral and
distal margins and about 7 non-plumose setae along mesial margin; endopod
with prominent basal apophysis, tip with 3 processes: caudal process broad-
ly rounded, forming a lateral shoulder; cannula short, cylindrical, endopo-
dial groove prominent; mesial process distally curved, rectangular, ob-
scuring cannula. Female pleopod 2 with 9-12 plumose setae along lateral
and distal margins, anterior surface with numerous setae.

Pleopod 3 about 2x as long as wide, distal segment about 1.6 length of
proximal segment, distal margin with about 20 long, plumose setae. Pleopod
4, exopod with setae along proximal lateral margin and single oblique sig-
moid suture. Pleopod 5 exopod with setae along proximal lateral margin,
and weak transverse suture.

Uropods of male very spatulate in mature specimens, slightly longer than
telson. Uropods of female cylindrical, equal or subequal to length of telson.

582 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

LY 7
a= LO
le

ie
|

Fig. 2. Caecidotea salemensis: a-f from male paratype, g from female paratype: a, Habitus,
dorsal; b, Antenna 2, proximal segments; c, Antenna 1; d, Same, distal segments; e, Mandibular
palp; f, Maxilla 1; g, Uropod, dorsal.

Etymology.—Named after the Salem Plateau.

Relationships.—Caecidotea salemensis is closest morphologically to C.
tridentata and two undescribed species from drain tiles in Missouri and
Illinois (Lewis and Bowman, in press), and Floyd County, Indiana (Lewis,
in prep.), referred to here as Caecidotea sp. #1 (Ill./Mo.) and Caecidotea
sp. #2 (Ind.). Caecidotea salemensis is readily separated from these species

VOLUME 94, NUMBER 2 583

Fig. 3. Caecidotea salemensis: a—d from male paratype, e from female paratype: a, Pereo-
pod 1; b, Pereopod 4; c, Same, dactyl; d, Pereopod 1, dactyl; e, Pereopod 1.

by the possession of a single sigmoid suture in pleopod 4; in the other three
Species two sutures are present. The palmar margins of the male pereopod
1 propus in C. salemensis and C. sp. #1 each bear one or two proximal
spines, a subtriangular median process and a bicuspid distal process. C.
tridentata differs in having a distinct fingerlike proximal process, while C.
sp. #2 has a shorter, triangular proximal process.

584 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

‘ \ f
X \ | 1 E pg Ty f
VK Hh Ap y
SHH! CU LOS
Bi A

KI
M
ih

Fig. 4. Caecidotea salemensis: a from female paratype, b—h from male paratype: a, Pleo-
pod 2; b, Pleopod 1; c, Pleopod 2; d, Same, tip of endopod, anterior; e, Same, posterior; f, Pleo-

pod 3; g, Pleopod 4; h, Pleopod 5.

VOLUME 94, NUMBER 2 585

The male pleopod 1 of all 4 species is similar in having long setae only
along the distal margin of the exopod. The male second pleopod endopod
tips are also very similar, although the placement of the endopodial groove
differs from one species to the next, and the relative sizes of the tip pro-
cesses also help to characterize the species. In C. salemensis, C. sp. #1
and C. sp. #2, the groove is on the lateral side of the mesial process, but
in C. tridentata it is on the mesial side of this process. The cannulas of C.
salemensis, C. tridentata and C. sp. #2 differ from that of C. sp. #1 in
being slender and low, mostly obscured from both the anterior and posterior
aspects. In C. sp. #1 the cannula protrudes beyond the mesial and caudal
processes, visible from both anterior and posterior aspects, and is recurved
mesially.

The low, partially obscured cannula, extending in a line parallel to the
axis of the endopod, plus the shape and setation of the male pleopod | of
Caecidotea salemensis, are typical of the morphology of the species of
Steeves’ (1964) Hobbsi Group. Within this group C. tridentata, C. sale-
mensis, C. sp. #1 and C. sp. #2 comprise an assemblage of closely related,
allopatric species.

Habitat and distribution.—Caecidotea salemensis is known from the
groundwater of caves, seeps, springs and cisterns. In caves it is usually
taken from streams and drip pools. These habitats are markedly different
from C. tridentata, C. sp. #1 and C. sp. #2, which are known only from
drain tiles and wells in areas where caves do not occur. Thus, C. salemensis
is a troglobite, whereas the other 3 species are phreatobites. C. salemensis
occurs widely in the Salem Plateau and is also known from a few other
localities near the edge of the plateau. Through much of its range C. sale-
mensis 18 Sympatric with C. antricola, a species which is distinct morpho-
logically from C. salemensis (Steeves, 1966; Lewis and Bowman, in press).

Remarks .—Ovigerous females were present in collections taken in March,
April and May with typically between 16 and 27 immatures present in the
brood pouches.

Caecidotea fustis, new species
Figs. 6, 7

Asellus stiladactylus.—Fleming, 1972:254 (in part).

Material examined (all deposited in the National Museum of Natural His-
tory, Smithsonian Institution)—MISSOURI: Crawford Co., Bat Cave, 7 mi
SE Leasburg, 16 Dec. 1973, JLC, 2 66; 24 May 1974, 2 64,4 2°; Bear
Cave, 8.5 mi NE Steelville, 6 Aug. 1980, JEG, 5 6¢, 11 22; Nameless
Cave, 16 Feb. 1974, JLC, 1 56, 1 2; Onondaga Cave, 4 mi SE Leasburg, 30
May 1974, JLC, 16,4 2 @; Pool in Onyx Cave, on bluff above Brazil Creek,

586 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Caecidotea salemensis: Variation of male pereopod | palmar margin (setae omitted):

a, Little Cave; b, Cistern, Imboden; c, Same; d, Cistern, Hughesville; e, Texas Cave; f, Lane
Cave (locality data in text).

3 mi N Campbell Bridge, 8 mi SE Bourbon, 20 July 1940, LH, 14 36, 4
2 2. Franklin Co., Copper Hollow Sink Cave, 8 Feb. 1958, O. Hawksley,
2566,6 29; Meramec Caverns (type-locality), drip pools, 2.5 mi SE Stan-
ton, 16 Dec. 1939, LH, 11.5 mm ¢ holotype, USNM 181301 and 13 ¢¢6
paratypes, USNM 181302; Seeps, 0.5 mi NW Mushroon Cave, 2.5 mi E
Sullivan, 5 Apr. 1942, LH, 32 ¢6, 22 2. Iron Co., seep, base of Taum
Sauk Mountain, near Mina Sauk Falls, 5 mi NW Hogan, 18 Jan. 1942, LH,
9 36,5 22. Washington Co., Corral Cave, 18 mi E Steelville, 21 Aug.
1980, JEG, 3 66,9 Qe.

Description.—Eyes vestigial or absent; pigment absent or lightly scattered
on dorsum. Longest male, 11.0 mm; body slender, linear, about 6.0 as
long as wide; coxae visible in dorsal view. Margins of head, pereonites and
telson moderately setose. Head about 1.3 x as wide as long, anterior margin
concave, postmandibular lobes slightly produced. Telson about 1.8 as long
as wide, sides subparallel, caudomedial lobe low, poorly defined.

Antenna | reaching to middle of distal segment of antenna 2 peduncle;
flagellum of about 13 segments, esthete formula 3-0-1-0-1 (Fig. 6b). Antenna
2 reaching about to anterior margin of telson; distal segment of peduncle
about 1.3 length of preceding segment; flagellum of about 63 segments.

Mandibles with 4-cuspate incisors and lacinia mobilis; spine row with
about 10 spines in left, 11 spines in right mandible. Palp with plumose setae
on distal segments. Maxilla 1, apex of outer lobe with 13 robust spines;
inner lobe with 5 apical plumose setae. Maxilliped with about 4 retinacula.

Male pereopod | propus about 1.5 as long as wide; palm with proximal
spine, triangular medial process and bicuspid distal process; dactyl flexor
margin with up to about 6 small spines, rounded process present in mature

VOLUME 94, NUMBER 2 587

Fig. 6. Caecidotea fustis: a~g from male paratype, Meramec Caverns; h, i from male from
Taum Sauk Mountain: a, Habitus, dorsal; b, Antenna 1, distal segments; c, Mandibular palp;
d, Pereopod 4; e, Maxilla 1, inner lobe; f, Same, outer lobe; g, Pereopod 1; h, Pereopod 1,
propus and dactyl, setae omitted; i, Uropod, dorsal.

specimens. Female pereopod | propus more slender, about 2x as long as
wide, palm with proximal spine, processes lacking; dactyl flexor margin
with about 3 spines. Pereopod 4 more robust in male than female.

Male pleopod 1 larger than pleopod 2; protopod about 0.6x length of
exopod, with 4—S retinacula; exopod about 2x as long as wide, with about

588 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 7. Caecidotea fustis: a from ovigerous female, Taum Sauk Mountain; b—h from male
paratype, Meramec Caverns: a, Pereopod 1; b, Pleopod 1; c, Pleopod 2; d, Same, tip of endo-
pod, anterior; e, Same, posterior; f, Pleopod 3; g, Pleopod 4; h, Pleopod 5.

5 plumose setae on distal margin and short setae on concave distal part of
lateral margin. Male pleopod 2 exopod, proximal segment with about 3
lateral setae, distal segment subtriangular, bearing about 13 plumose setae
along lateral and distal margins; endopod with prominent basal apophysis,

VOLUME 94, NUMBER 2 589

tip with 4 processes: (1) cannula obscured by other processes, forming low
truncate conical extension of endopodial groove; (2) mesial process broad,
suboval, distally rounded; (3) lateral process high, digitiform, slightly re-
curved laterally, with low subtriangular flange on caudal surface; and (4)
caudal process forming heavily sclerotized band, broadening laterally,
across posterior distal part of endopod.

Pleopod 3 about 2.1 as long as wide, distal segment about 1.3x length
of proximal segment, distal margin with about 6-7 long, plumose setae.
Pleopod 4, exopod with setae along proximal lateral margin, single oblique
sigmoid suture present. Pleopod 5 with proximal seta, 2 weak sutures.

Uropods of male spatulate, about 1.7x< length of telson, exopod distinctly
club-shaped in mature males. Uropods of female cylindrical, equal or sub-
equal to length of telson.

Etymology.—The name, proposed as a noun, is derived from the Latin
‘‘fustis’’? (=club), referring to the appearance of the uropods of mature
males.

Relationships.—The general morphological affinities of Caecidotea fustis
are with the members of the Hobbsi Group, sharing with other members of
this group the shape and distal setation of the male pleopod 1, and the low,
truncate distally extending cannula. Specifically, C. fustis may be related
to C. stiladactyla. The male pereopod 1 of these species is quite similar in
most specimens, although in large specimens of C. fustis the proximal spine
present on the palmar margin of the propus in both species is replaced by
a large, triangular process. The endopod tips of C. fustis and C. stiladactyla
are similar in that each possesses a truncate, conical cannula which is nearly
obscured by other, somewhat digitiform processes. However, C. stiladac-
tyla exhibits a phenomenon which has not been reported to occur in other
asellids, i.e., the cannula has both recumbent and erect positions, and ap-
parently is capable of a telescoping motion within the tip of the endopod.
Fleming (1972) illustrated both positions, but the recumbent position ap-
pears to be the endopod tip of C. fustis (reported from Onyx Cave, Missouri
by Fleming) rather than that of C. stiladactyla. Besides the lack of the tele-
scoping cannula, C. fustis is pigmented in some collections, while pigmen-
tation is unknown in C. stiladactyla, and the male pleopod 1 of C. fustis
bears distal plumose setae which are absent in C. stiladactyla.

Habitat and distribution.—Caecidotea fustis is known only from caves
and seeps in a narrow north-south band lying between Washington and Iron
counties, Missouri. This area, on the western flank of the St. Francois
Mountains, is within the northeastern part of the Salem Plateau. Caecidotea
fustis is allopatric with C. stiladactyla, which is apparently known authenti-
cally only from localities in Arkansas which are adjacent to the edge of the
Salem Plateau.

590 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Acknowledgments

I would like to thank Dr. Thomas E. Bowman for both reading the manu-
script and expediting the loan of material from the collections of the National
Museum of Natural History, including all of the specimens donated to the
museum by Mr. Leslie Hubricht. Mr. James E. Gardner donated many
specimens collected during his work on a cooperative cave inventory by the
Missouri Department of Conservation, the Mark Twain National Forest,
and the North Central Forest Experiment Station, and provided field assis-
tance to me on a collecting trip through Missouri in August 1979. Mr. John
L. Craig also donated his large collection of Missouri cave asellids, which
added many localities for the new species described herein. Dr. Stuart Neff
both read the manuscript and made travel funds available through the Water
Resources Laboratory, University of Louisville. The Research Advisory
Committee of the National Speleological Society provided grant funds par-
tially used for travel expenses incurred during a collecting trip through the
Ozarks. Finally, Mrs. Teresa M. Lewis provided field assistance and moral
support during the preparation of this paper.

Literature Cited

Bretz, J. H. 1956. Caves of Missouri.—Missouri Division of Geological Survey and Water

Resources, volume 39, 2nd series, 490 pp.

, and S. Harris. 1961. Caves of Illinois.—Illinois State Geologic Survey, Report of

Investigations 215, 87 pp.

Craig, J. L. 1977. Invertebrate faunas of caves to be inundated by the Meramec Park Lake
in eastern Missouri.—National Speleological Society Bulletin 39(3):80-89.

Creaser, E. P. 1931. A new blind isopod of the genus Caecidotea from a Missouri cave.—
Occasional Papers of the Museum of Zoology, University of Michigan 222: 1-7.
Fleming, L. E. 1972. The evolution of the eastern North American isopods of the genus
Asellus (Crustacea: Asellidae).—International Journal of Speleology 4:221-256.

Lewis, J. J., and T. E. Bowman. The subterranean asellids of Illinois.—Smithsonian Contri-
butions to Zoology 335. [in press. ]

Mackin, J. G., and L. Hubricht. 1940. Descriptions of seven new species of Caecidotea
(Isopoda: Asellidae) from the central United States.—Transactions of the American
Microscopial Society 59(3):383-397.

Steeves, H. R. 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.

Vineyard, J. D., and G. L. Feder. 1974. Springs of Missouri.—Missouri Geological Survey
and Water Resources, Report 29, 266 pp.

Department of Biology, University of Louisville, Louisville, Kentucky
40208.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 591-597

A REVIEW OF THE PLATYSQUILLA COMPLEX
(CRUSTACEA, STOMATOPODA, NANNOSQUILLIDAE),
WITH THE DESIGNATION OF TWO NEW GENERA

Raymond B. Manning and David K. Camp

Abstract.—Platysquilla Manning, 1967, is redefined, with P. eusebia
(Risso, 1816) as its only species. The other species previously assigned to
Platysquilla, Lysiosquilla enodis Manning, 1962, and Platysquilla horologii
Camp, 1971, are removed and made the type-species of two new, monotypic
genera, Platysquilloides and Mexisquilla, respectively. The three genera
are defined, and the type-species are re-illustrated.

The nomenclatural histories of Squilla eusebia Risso, 1816, and Lysios-
quilla enodis Manning, 1962, were briefly summarized by Camp (1971:21).
Both had been transferred to Heterosquilla (Manning, 1963) and later to
Platysquilla (Manning, 1967), with S. eusebia designated as type-species of
the latter genus.

When Platysquilla horologii Camp, 1971, was described, the three known
specimens did not fully agree with the definition of the genus, although it
was recognized that they agreed more closely with Platysquilla than with
any other established genus. Therefore, a very conservative approach was
taken, and the generic description was emended to allow the inclusion of
P. horologii in Platysquilla. With the discovery of additional specimens of
P. horologii (R. W. Heard and D. K. Camp, in prep.), and after further
consideration of the relationships between the three species, we believe the
latter are sufficiently different from one another to warrant their separation
into three monotypic genera.

Platysquilla Manning, 1967

Platysquilla Manning, 1967:238.—Holthuis, 1967:26.—Manning, 1968:111;
1969:90.—Camp, 1971:120.—Manning, 1977:93; 1980:368 [In part].

Definition.—Size moderate, total length 75 mm or less; body smooth,
depressed, loosely articulated; eye of moderate size, cornea expanded but
not bilobed, set obliquely on stalk; rostral plate subquadrate with apical
spine, lateral margins straight, anterolateral corners rounded; antennal pro-
topod with 2 mesial and 2 ventral papillae; carapace narrowed anteriorly,
without carinae or spines, cervical groove indicated on lateral plate only;
thoracic somites without dorsal carinae, lateral margins truncate; eighth —

592 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Platysquilla eusebia, female, 44.5 mm, Naples, Italy, USNM 23207: a, Front; b,
Claw; c, Sixth abdominal somite, telson, and uropod; d, Telson, ventral view; e, Uropod,
ventral view.

thoracic somite without low, inconspicuous tubercle on midline of ventral
surface; 5 epipods present; mandibular palp absent; raptorial claw slender,
dactylus not inflated basally; outer margin of dactylus faintly notched ba-
sally; propodus fully pectinate with 4 movable spines at base, first longest,
second shortest; carpus with distal, dorsal spine; ischiomeral articulation
terminal; merus slender, elongate, longer than ischium; endopods of walking
legs 2-segmented, distal segment of first 2 legs ovate, that of last leg slen-
derest; abdomen depressed, loosely articulated, anterolateral plates with
complete suture; sixth abdominal somite without posterolateral spines, with
curved, ventrally directed process present on each side in front of articu-
lation of uropod, with 2 strong spines on ventral, posterior margin; telson
broad, with obtuse, triangular, wide median posterior projection; marginal
armature of telson consisting of row of slender submedian denticles, 1 mov-
able submedian tooth, and 4 sharp, fixed lateral teeth, usually with slender
denticle or tubercle between fixed teeth; basal segment of uropod with 2
dorsal carinae, inner terminating in slender spine; proximal segment of ex-

VOLUME 94. NUMBER 2 593

opod with short dorsal carina, outer margin armed with slender, movable,
sharp spines; endopod triangular, with proximal portion of outer edge folded
over dorsally; spines of basal prolongation triangular in cross-section, inner
spine longer. [Modified from Manning, 1969:90. ]

Type-species.—Squilla eusebia Risso, 1816, by original designation. See
Figure 1.

Gender.—Feminine.

Number of species.—One.

Distribution.—Mediterranean Sea and adjacent North Atlantic, off Por-
tugal, France, and the west coast of Ireland (Manning, 1977).

Platysquilloides, new genus

Platysquilla Manning, 1967:238.—Holthuis, 1967:26.—Manning, 1968:111;
1969:90.—Camp, 1971:120.—Manning, 1977:93; 1980:368 [In part].

Definition.—Size moderate, total length 70 mm or less; body smooth,
depressed, loosely articulated; eye of moderate size, cornea faintly bilobed,
set obliquely on stalk; rostral plate subquadrate with apical spine, lateral
margins slightly convex, anterolateral corners rounded; antennal protopod
with 1 mesial and 2 ventral papillae; carapace narrowed anteriorly, without
Carinae or spines, cervical groove indicated on lateral plate only; thoracic
somites without dorsal carinae, lateral margins truncate; eighth thoracic
somite with low, inconspicuous tubercle on midline of ventral surface; 4
epipods present; mandibular palp absent; raptorial claw slender, dactylus
not inflated basally; outer margin of dactylus strongly notched basally; pro-
podus fully pectinate, with 4 movable spines at base, first longest, second
shortest; carpus with distal, dorsal spine; ischiomeral articulation terminal;
merus slender, elongate, longer than ischium; endopods of walking legs 2-
segmented, distal segment of first 2 legs ovate, that of last leg slenderest;
abdomen depressed, loosely articulated, anterolateral plates with complete
suture; sixth abdominal somite with sharp posterolateral spines, with
curved, ventrally directed process present on each side in front of articu-
lation of uropod, without spines on ventral, posterior margin; telson broad,
with obtuse, subtriangular, narrow median posterior projection; marginal
armature of telson consisting of row of slender submedian denticles, 1 mov-
able submedian tooth, and 4 fixed lateral teeth, mesial 2 spatulate, with
slender denticle between fixed teeth; basal segment of uropod with 2 dorsal
Carinae, inner terminating in slender spine; proximal segment of exopod
with short dorsal carina, outer margin armed with spatulate, movable spines;
endopod triangular, with proximal portion of outer edge folded over dor-
sally; spines of basal prolongation triangular in cross-section, inner spine
longer.

Type-species.—Lysiosquilla enodis Manning, 1962. See Figure 2.

594 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Platysquilloides enodis, male, 48 mm, off New Jersey, USNM 172218: a, Front;
b, Claw; c, Sixth abdominal somite, telson, and uropod; d, Telson, ventral view; e, Uropod,
ventral view.

Etymology.—The name is composed of the generic name Platysquilla
with the Latin suffix **-oides,’’ meaning ‘‘resembling.”’

Gender.—Masculine.

Number of species.—One.

Remarks.—This new genus differs from Platysquilla in having 4, rather
than 5, epipods and in having | mesial papilla on the antennal peduncle,
rather than having 2 mesial papillae. It shares other, less significant, char-
acteristics with the other western Atlantic genus described below which
further separate them from the eastern Atlantic Platysquilla. These include
the presence of a strong, rather than faint, proximal notch on the dactylus
of the raptorial claw, the lack of submedian spines on the ventral, posterior
margin of the sixth abdominal somite, the presence of posterolateral spines
on the sixth abdominal somite, a narrow, rather than broad, posterior me-
dian projection on the dorsum of the telson, and spatulate, rather than sharp,
movable spines on the outer margin of the exopod of the uropod.

Distribution.—Northwestern Atlantic, from scattered localities between
Massachusetts and North Carolina (Manning, 1969), including 16 km N of
Atlantic City, New Jersey, and off Maryland and Virginia (Howells, Karp,
and Langton, 1980).

VOLUME 94, NUMBER 2 ; 595

Fig. 3. Mexisquilla horologii, female paratype, ca. 16 mm, west of Egmont Key, Florida,
USNM 128832: a, Front; b, Claw; c, Sixth abdominal somite, telson, and uropod; d, Telson,
ventral view; e, Uropod, ventral view.

Mexisquilla, new genus
Platysquilla.—Camp, 1971:120 [In part].

Definition.—Size small, total length 20 mm or less; body smooth, de-
pressed, loosely articulated; eye of moderate size, cornea faintly bilobed,
set obliquely on stalk; rostral plate cordiform, lateral and anterolateral mar-
gins broadly rounded, sloping to obtuse apex; antennal protopod without
mesial papillae, with 2 ventral papillae; carapace narrowed anteriorly, with-
out carina or spines, cervical groove indicated on lateral plate only; thoracic
somites without dorsal carinae, lateral margins truncate; eighth thoracic
somite with low, inconspicuous tubercle on midline of ventral surface; 3
epipods present; mandibular palp absent; raptorial claw slender, dactylus
not inflated basally; outer margin of dactylus strongly notched basally; pro-
podus fully pectinate, with 4 movable spines at base, first longest, second
shortest; carpus with distal, dorsal spine; ischiomeral articulation terminal;
merus slender, elongate, longer than ischium; endopods of walking legs 2-

596 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

segmented, distal segment of first 2 legs ovate, that of last leg slenderest;
abdomen depressed, loosely articulated, anterolateral plates with complete
suture; sixth abdominal somite with sharp posterolateral spines, with
curved, ventrally directed process present on each side in front of articu-
lation of uropod, without spines on ventral, posterior margin; telson broad,
with obtuse, subtriangular, narrow median posterior projection; marginal
armature of telson consisting of row of slender submedian denticles, 1 mov-
able submedian tooth, and 4 fixed lateral teeth, mesial 2 sharp, with slender
denticle between fixed teeth; basal segment of uropod with 2 dorsal carinae,
inner terminating in slender spine; proximal segment of exopod with short
dorsal carina, outer margin armed with spatulate, movable spines; endopod
triangular, with proximal portion of outer edge folded over dorsally; spines
of basal prolongation triangular in cross-section, inner spine longer.

Type-species.—Platysquilla horologii Camp, 1971. See Figure 3.

Etymology.—Mexisquilla is derived from Gulf of Mexico and the generic
name Squilla. It alludes to the apparent restriction of members of the genus
to the Gulf of Mexico.

Gender.—Feminine.

Number of species.—One.

Remarks.—This new genus differs from Platysquilla in having 3, rather
than 5, epipods and in having no mesial papillae on the antennal peduncle
rather than having 2 such papillae. Other characteristics which differ from
those of Platysquilla, but which are shared with Platysquilloides, are dis-
cussed under the latter genus.

Mexisquilla also differs from Platysquilloides in having fewer epipods and
fewer antennal papillae than are present on the latter genus.

Mexisquilla differs from both Platysquilla and Platysquilloides in its
overall size and degree of development. Mexisquilla horologii is relatively
much smaller than the type-species of the other two genera. It also shows
reduction in development of the telson characters relative to such devel-
opment in Platysquilla eusebia and Platysquilloides enodis. The possibility
of neotenic origin of M. horologii was discussed by Camp (1973:18).

Distribution.—Eastern Gulf of Mexico (Camp, 1971, 1973).

Acknowledgments

We thank F. A. Chace, Jr., L. B. Holthuis, and W. G. Lyons for com-
menting on the manuscript, Lilly King Manning for preparing the illustra-
tions, and R. W. Heard for offering specimens of M. horologii for study.

Literature Cited

Camp, D. K. 1971. Platysquilla horologii (Stomatopoda, Lysiosquillidae), a new species from
the Gulf of Mexico, with an emendation of the generic definition.—Proceedings of the
Biological Society of Washington 84(15):119-128.

VOLUME 94, NUMBER 2 597

. 1973. Stomatopod Crustacea.—Memoirs of the Hourglass Cruises 3(2):1—100.

Holthuis, L. B. 1967. Fam. Lysiosquillidae et Bathysquillidae. In H. E. Gruner and L. B.
Holthuis, eds., Crustaceorum Catalogus 1:1—28. W. Junk, The Hague.

Howells, R. G., C. Karp, and R. W. Langton. 1980. Occurrence of a rare mantis shrimp,
Platysquilla enodis (Manning, 1962), in the Middle Atlantic Bight area (Stomatopoda).—
Crustaceana 38(1):101-104.

Manning, R. B. 1962. Seven new species of stomatopod crustaceans from the northwestern

Atlantic.—Proceedings of the Biological Society of Washington 75:215—222.

. 1963. Preliminary revision of the genera Pseudosquilla and Lysiosquilla with descrip-

tions of six new genera (Crustacea: Stomatopoda).—Bulletin of Marine Science of the

Gulf and Caribbean 13(2):308-328.

. 1967. Preliminary account of a new genus and a new family of Stomatopoda.—Crus-

taceana 13(2):238-239.

. 1968. A revision of the family Squillidae (Crustacea, Stomatopoda), with the descrip-

tion of eight new genera.—Bulletin of Marine Science 18(1):105—142.

1969. Stomatopod Crustacea of the western Atlantic.—Studies in Tropical Ocean-
ography, No. 8: viii + 380. Institute of Marine Sciences, University of Miami.

. 1977. A monograph of the West African stomatopod Crustacea.—Atlantide Report

No. 12:25-181.

. 1980. The superfamilies, families, and genera of Recent stomatopod Crustacea, with

diagnoses of six new families.—Proceedings of the Biological Society of Washington

93(2):362-372.

Risso, A. 1816. Histoire naturelle des Crustacés des environs de Nice, pp. 1-175, 3 pls. Paris.

(RBM) Department of Invertebrate Zoology, National Museum of Natural
History, Smithsonian Institution, Washington, D.C. 20560; (DKC) Florida
Department of Natural Resources, Marine Research Laboratory, 100 Eighth
Avenue SE, St. Petersburg, Florida 33701.

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 598-621

ANATOMY OF DIASTOMA MELANIOIDES (REEVE,
1849) WITH REMARKS ON THE SYSTEMATIC
POSITION OF THE FAMILY DIASTOMATIDAE
(PROSOBRANCHIA: GASTROPODA)

Richard S. Houbrick

Abstract.—Diastoma melanioides is the living survivor of a long lineage
of snails in the Family Diastomatidae that occurred in the Tethys Sea during
the Tertiary. Study of the anatomy shows that this species has open pallial
gonoducts, and aphallic males, which establishes it in the superfamily Cer-
ithiacea. A large ovipositor on the right side of the foot extends into the
mesopodium. The radula is taenioglossate; the alimentary tract has paired
salivary glands that run through the nerve ring and a stomach with style
sac, gastric shield and reduced spiral caecum. The cephalic cavity is large
and accommodates a small buccal mass and lengthy buccal and labial
nerves. The nervous system is moderately condensed. Living animals occur
subtidally in shallow water where they burrow in sandy bottoms and graze
on algae and detritus. Development appears to be direct. Diastoma mela-
nioides most closely resembles members of the Cerithiidae in anatomy and
ecology. Anatomical and shell characters, and the fossil record indicate that
Diastoma should be given familial status. The family is placed close to the
Cerithiidae, the Potamididae, and the freshwater Melanopsidae.

Introduction

Diastoma melanioides (Reeve) is a relatively unknown cerithiacean
prosobranch of moderate size which is restricted to a limited area of the
coastline of the Great Australian Bight. It is the sole survivor of the Diasto-
ma lineage, Family Diastomatidae. The taxonomic limits of this family have
been poorly defined; consequently, a number of heterogenous groups have
been referred to the Diastomatidae by numerous authors, and the family
has been expanded well beyond the original concept. The living relict, Dias-
toma melanioides, heretofore known only from its shell, is rare in most
museum collections and unfigured in most publications. Recently, some
well-preserved specimens collected at Esperance, Western Australia, were
sent to me through the kindness of Mr. Alan Longbottom. This material has
provided information about the operculum, radula, and anatomy of Diasto-
ma melanioides which unequivocally establishes that it is a cerithiacean.

The account that follows presents a description of the shell and internal

VOLUME 94, NUMBER 2 599

anatomy, reviews the literature, and sets definite systematic limits to the
family by providing familial and generic descriptions and a synonymy.

Material arid Methods

Fourteen specimens of Diastoma melanioides were sent to me for ex-
amination by Mr. Alan Longbottom of Esperance, Western Australia. Eight
of these were well-preserved animals and were dissected under a Wild M-5
stereo dissecting microscope. A one percent Methylene Blue aqueous so-
lution was used as a stain. Protoconchs, shell ultrastructure, radula and
periostracum were studied with a Nova-Scan SEM. All measurements are
relative to average-sized snails (see Table 1). There was not sufficient ma-
terial for statistical measurements of the soft parts.

Although anatomical observations are substantially accurate, the limited
material and its preserved state may not have shown all the details of the
fragile pallial gonoduct, stomach and smaller nerves. My work should be
reconfirmed by the study of living snails.

Description

List of abbreviations.—a—anus; bg—buccal ganglion; bm—buccal
mass; bt—buccal tensor; cbhc—cerebral-buccal connective; cc—cerebral
cavity; cg—cerebral ganglion; cme—cut mantle edge; ct—ctenidium; d—
dialyneury; dg—digestive gland; d/n—dorsal labial nerve; es—esophagus;
exh—exhalant siphon; f—foot; ff—fold emerging from spiral calcum; ga—
glandular area; gs—gastric shield; hg—hypobranchial gland; i—intestine;
i/—inner lamina; inh—inhalant siphon; j—jaws; k—kidney; //n—lateral la-
bial nerve; /mn—left mantle nerve; /pg—left pleural ganglion; /sg—left sal-
ivary gland; m—mouth; mn—mantle nerve; mp—mantle papillae; mpg—
mesopodial ganglion; ng—nerve ganglion; nr—nerve ring; od—oviduct;
odd—opening of digestive diverticula; og—oviducal groove; o/—outer lam-
ina; on—optic nerve; op—operculum; ovp—ovipositor; os—osphradium;
osd—opening of salivary duct; ov—ovary; pes—posterior esophagus; pg—
pedal ganglion; pod—proximal end of pallial oviduct; ppg—propodial
groove; psg—proximal end of salivary gland; r—rectum; rd—radula; rcg—
right cerebral ganglion;-rpg—tright pleural ganglion; rsg—right salivary
gland; sa—sorting area; sbg—subesophageal ganglion; sbn—subvisceral
nerve; sby—subvisceral connective; sc—spiral caecum; sd—sperm duct;
sec—supraesophageal connective; sg—salivary gland; sm—snout; spg—su-
praesophageal ganglion; sp—spermatophore receptacle; sr—seminal recep-
tacle; ss—style sac; st—stomach; t/—typhlosole 1; t2—typhlosole 2; tn—
tentacle nerve; wcc—wall of cerebral cavity.

Specimens examined.—Abbreviations: AMS = Australian Museum,

600 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Sydney; USNM = United States National Museum; WAM = Western Aus-
tralian Museum.

Western Australia: Cheyne Beach, 40 mi E of Albany, 34°47’S, 118°25’E
(AMS); Cheyne Beach, near Posidonia, 34°47’S, 118°25’E (WAM); Cape
Riche, 70 mi E of Albany, 34°37’S, 118°46’E (AMS); North Side of High
Island, Duke of Orleans Bay, 33°55’S, 122°37'E (WAM); Esperance Bay,
Esperance (USNM, 806583, 801614); Mississippi Bay, 30 mi E of Esperance,
34°00'S, 122°17’E (AMS); Eyre Highway, 65 mi E of Madura, 31°55’S,
127°00'E (AMS).

South Australia: Port Sinclair, 32°06’S, 133°00' E (AMS); Smokey Bay,
32°20'S, 133°47'E (AMS); Streaky Bay, 32°32’S, 134°08’E (AMS).

Shell (Table 1; Fig. 1A—D, G—J).—Shell elongate, length 30-50 mm, tur-
reted, having apical angle of 25 degrees and comprising 10-13 convex
whorls. Protoconch (Fig. 11) has two convex, smooth whorls. Transition
from protoconch to teleoconch sharply defined by straight lip of protoconch
and sudden change in sculpture. First two whorls of teleoconch sculptured
with 3 weak spiral cords which increase to 4 on the third whorl. Spiral cords
weak, increasing in number but diminishing in intensity on each successive
whorl. Slanting axial ribs appear on fourth teleoconch whorl and are most
pronounced and numerous (12-18) on median whorls of shell but become
weaker and nearly lost on penultimate and body whorls. Overall fine can-
cellate sculpture results from intersection of axial ribs and spiral cords.
Varices occasionally present as low flat axial ridges. Suture straight, deeply
incised, forming ramp at posterior of each whorl that becomes most pro-
nounced on body whorl where outer lip of aperture is slightly detached from
body whorl to form anterior apertural notch. Aperture (Fig. 1H) tear-shaped,
wide at base and a little over one-third the length of shell. Anterior siphonal
canal virtually absent but distinguishable as shallow, depressed curve of
apertural basal lip. No anal canal present. Columella slightly convex with
. Slight but distinct median oblique fold originating from base of outer aper-
tural lip and continuing into aperture of the shell where it ends about halfway
up body whorl. Slight columellar callus present. Outer lip of aperture con-
tinuous, not broken by anterior canal, thin, smooth and slightly curved.
Shell color white, flecked with small, spirally arranged, light brown spots
and blotches. Largest blotches at posterior of each whorl, adjacent to su-
ture.

_ Periostracum (Fig. 3D—F) thin and tan colored, covering entire shell and

with spiral rows of tiny hair-like projections that correspond to underlying
spiral sculpture of shell. Periostracum on live collected specimens has over-
all fuzzy appearance but flakes off easily when dry.

Operculum (Fig. 1J, K) dark brown, ovate and paucispiral with nucleus near
the columellar edge. Operculum closes aperture when animal withdrawn.

VOLUME 94, NUMBER 2 601

Fig. 1. A—D. Dorsal, ventral and side views of the shell of Diastoma melanioides (49.7
mm long). Shell in A has been whitened with Ammonium chloride to enhance shell sculpture;
E-F, Shells of the Eocene fossil Diastoma costellata, from the Paris Basin. Note the pro-
nounced sutural ramp at aperture. (52 mm long). G—H. Shell apex (G), and adult whorls and
aperture (H) of Diastoma melanioides showing details of shell sculpture; I, SEM micrograph
of protoconch of Diastoma melanioides showing change from embryonic to juvenile shell
sculpture; J-K. Operculum showing free surface (J) and attached side with muscle scar (K).

Animal: external features (Fig. 2A, B, D).—Base color white to flesh with
a few tiny flecks of brownish pigment on head-snout. Head with a large,
highly extensible, spade-shaped snout (sn) deeply incised with transverse
wrinkles. Cephalic tentacles short, about one-third the length of snout, and
widening to a peduncle where attached to head. A tiny black eye present

602 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Analysis of shell characters of Diastoma melanioides. (Measurements in mm.)

Character (n = 10) x SD Range
Length 38.58 5.93 3049.68
Width 11.73 1.34 9.48-14.08
No. of whorls We 7/ 0.95 10-13
No. axial ribs per whorl 16.1 1.97 12-18
Aperture length 11.06 1.28 9.25-13.44
Aperture width 4.87 0.45 4.20-5.82

on outer edge of each peduncular stalk. The relatively large foot (f/) has a
propodial grove (ppg) (anterior pedal mucous gland). A furrow emerges
from the exhalant siphon and runs down the right side of the foot. In fe-
males, a large, bulbous, vermiform ovipositor emerging from a deep pit is
present on the median right side of the foot (Fig. 2B, C). Mantle edge thick,
characterized by a sharply defined ridge at the exhalant siphon (exh). Upper
margin of mantle edge bifurcate, with thin, membranous fringe from which
emerges another papillated fringe (mp). Mantle papillae are short and fused
with each other at their bases. Inhalant siphon (inh) marked by a thickening
of mantle wall and by papillae wider than those at dorsal mantle edge. No
papillae on ventral edge of mantle. Mantle thin, unpigmented, forming dorsal
fold that begins at exhalant siphon and extends back for one-half of the first
whorl. Major mantle organs visible through mantle wall. Tan colored, single-
lobed kidney (k) of moderate size present. Digestive gland (dg) dark brown.
Gonads occupy dorsal parts of upper whorls.

Mantle cavity and associated organs (Fig. 2D).—The deep, spacious man-
tle cavity occupies the last two whorls of the animal. A thick glandular area
(ga) lies on the inner side of the mantle edge adjacent to the tip of the
inhalant siphon. The bipectinate osphradium (os) is a thin brown ridge com-
prising about 140 thick filaments and is about 13.5 mm long and 0.2 mm
wide. It lies adjacent to the ctenidium but is separated from it by a narrow
margin of thin mantle tissue. The osphradium curves away from the ctenid-
ium at its distal end and twists toward the inhalant siphon. The monopec-
tinate ctenidium (cf) is large, about 17 mm long, 2.3 mm wide, and comprises
195 thin leaflets. A long, wide hypobranchial gland (hg) lies adjacent to the
ctenidium and excretes much mucus. It is composed of thick transverse
folds which become darker and wider at the distal end, adjacent to the anus
and exhalant siphon. The osphradium, ctenidium and hypobranchial glands
do not extend the length of the mantle cavity but end about a third of the
way from its proximal end. The rectum (r) is filled with ovoid fecal pellets
comprised of sand grains, detritus, and algal fragments. The pallial gono-
ducts extend the length of the mantle cavity, and are open in both sexes. The

VOLUME 94, NUMBER 2 603

sn OH D
Fig. 2. A, Diastoma melanioides, female removed from shell and viewed from right dorsal
side; B, Female showing large bulbous ovipositor on right side of foot and ciliated groove
leading to it from exhalant siphon; C, Midsagittal section through head-foot showing interior
of ovipositor with mucus gland, central cavity and innervation by nervous system; D, Female
removed from shell, mantle cavity opened along left side and mantle folded to right.

spacious proximal mantle cavity contains a large pericardial sac that accom-
modates a large auricle and ventricle. The single-lobed kidney (Fig. 2A, k)
has a large slit-like opening on its ventral surface that leads into the mantle
cavity.

604 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Alimentary system (Fig. 4).—Diastoma has a spacious cephalic cavity
(hemocoel). The highly extensible snout is dominated by bundles of circular
muscle. The mouth (m) is deeply recessed between the two lobes of the
snout apex. The jaws (j) are thin and brittle, each about 1.3 mm long. Under
the microscope, they appear scale-like near their cutting edges. The radular
ribbon (Fig. 3A—C) is tiny, about 3.8 mm long, 0.6 mm wide and comprising
50 rows of teeth. It is about one-eleventh the length of the shell and is
typically taenioglossate (2 + 1 + 1+ 1 + 2), not unlike those of some Cer-
ithium species. Rachidian tooth quadrate in shape, concave laterally and
convex basally. Cutting edge of rachidian has spade-shaped, pointed, central
cusp flanked on each side with 2—4 smaller blunt denticles. Basal plate of
rachidian tooth has pair of basolateral projections. Lateral tooth rhombiodal
in shape, laterally elongate where it is attached onto the basal radular mem-
brane. Top of lateral tooth has cutting edge with tiny inner denticle, a large,
triangular, pointed cusp and 2-3 tiny denticles, respectively. Base of lateral
tooth straight. Basal plate with centrally located, small, blunt peg. Marginal
teeth long, slender, swollen centrally, curving and with spatulate tips. Cut-
ting edge of inner marginal tooth has 2 sharp inner denticles, a long, pointed,
central cusp and a short blunt outer denticle. Outer marginal tooth identical
but lacks outer denticle.

The buccal mass (bm) is relatively small, about 3 mm long, and is attached
anteriorly to the inner tip of the cephalic cavity by numerous, thin muscular
tensors (bt). Originating at the center of the ventral buccal mass is a long,
post-median retractor muscle that inserts on the ventral wall of the cephalic
cavity. It is flanked by 2 smaller retractors. The radula sac originates at the
central posterior part of the buccal mass and extends dorsally.

The esophagus (es) is long, narrow and loosely connected to the base and
sides of the buccal body cavity by several long, thin muscles. The largest
of these extend from the esophagus to the nerve ring and then to the walls
of the cephalic cavity (m). The esophagus may thus be partially pulled
through the nerve ring when the snout is extended. Attached to the dorsal
surface of the anterior esophagus are a pair of long tubular, convoluted
salivary glands (sg) which pass through the nerve ring and end on the mid-
esophagus, where they are twisted to its left side (Fig. 5D). Each salivary
gland empties into a side of the median buccal cavity (Fig. 4, osd). The

as

Fig. 3. SEM micrographs: A, Radula of Diastoma melanioides showing unworn portion of
radular ribbon (0.6 mm wide); B, Details of rachidian, lateral and marginal teeth (0.3 mm wide);
C, Close up of lateral tooth (0.15 mm long) showing its insertion on basal radular membrane;
D, SEM micrograph of protoconch and juvenile whorls of shell to show early sculpture and
periostracum (15x); E, SEM of periostracum illustrating microscopic periostracal hairs (50x);
F, Cross section of shell showing periostracum at top and lamellar aragonite below (50x).

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606

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

WCC
cbc
rsg

lin

tn

rcg
rpg

lmn

‘PSg
sbn

pes

Fig. 4. Dissection of head of Diastoma melanioides opened by a dorsal longitudinal cut to
expose cephalic cavity and anterior alimentary tract (buccal mass 3 mm long).

VOLUME 94, NUMBER 2 607

anterior esophagus has a dorsal food groove and two dorsal ridges (Fig. 5E)
which become ventral-lateral (Fig. 5F) as they pass through the nerve ring.
The mid-esophagus is flattened dorso-ventrally and the dorsal food ridges
become more laterally placed.

The stomach (Fig. 5G) is typically cerithiacean and is a large organ almost
one and a half whorls in length. It has a well-defined sorting area (sa)
comprised of transverse lamellae-like folds, a style sac (ss), a large cuticular
gastric shield (gs) and 2 liver ducts (odd). Posterior to the gastric shield is
a large pad-like area (ff) of uncertain function which arises from the reduced
spiral caecum (sc). No crystalline style was found but this was probably
due to the preserved state of the animals.

Reproductive system (Fig. 5A).—The presence of open pallial gonoducts,
aphallic males, spermatophores, and the location of the seminal receptacle
are typically cerithiacean. Males are smaller than females and easily distin-
guished from them by the lack of an ovipositor on the right side of the foot.
The riarrow laminae of the pallial gonoducts are thin walled and glandular
at the proximal end of the gonoduct where the outer lamina becomes thick-
ened and white along its base. This area is probably the prostate and sper-
matophore-forming organ. The white testis overlays the digestive gland on
the dorsal surface of each upper whorl.

The long pallial gonoduct of females is a larger, wider organ than in males
(Figs. 2D, 5A). The deep slit-like oviducal groove (og) formed by the 2
laminae of the open duct is lined with numerous transverse folds of glandular
tissue which become thick and more opaque at the proximal end of the duct.
This is the albumen gland (ag). The pallial oviduct ends just anterior to the
anus. A narrow slit, the sperm-collecting gutter (sd), lies along the free edge
of the outer lamina (o/) enlarging to form a pouch-like seminal receptacle
(sr) which is located in the post-median section of the lamina. The seminal
receptacle is a flattened, kidney-shaped structure with a spongy interior that
initially receives the spermatophore and also serves to hold the sperm. A
central axial flap of tissue divides it into 2 chambers that join at the end. I
extracted the remains of what appeared to be a disintegrating spermatophore
from the inner chamber. The seminal receptacle is narrow at its proximal
end and becomes a blind tube (sr) in the proximal part (pod) of the outer
lamina of the oviduct. No opening from the receptacle to the oviducal
groove was found.

The ovary (Fig. 2A, ov) lies on the dorsal surface of the digestive gland
(dg) and is light tan. Oocytes are about 0.4 mm in diameter and are shaped
like rounded triangles.

One of the most unusual structures associated with the female reproduc-
tive tract of Diastoma is the large ovipositor (Fig. 2B, C) on the right median
part of the mesopodium. This ball-like pad has a crescent-spiral configura-
tion that sinks into a deep pit at its center (Fig. 2C, ovp). The pit leads into

608 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. A, Diagrammatic representation of pallial oviduct with sections of duct cut away
to show arrangement of laminae and associated ducts, spermatophore and seminal receptacles.
Pallial oviduct is 10 mm long and oriented with distal end at bottom of figure; B, Lateral view
of cerebropedal complex; C, Dorsal view of nerve ring and associated nerves; D, Anterior and
mid-esophagus with nerve ring and salivary glands; E, Section through anterior esophagus
showing dorsal folds and salivary ducts; F, Section through mid-esophagus showing ventral-
lateral (dorsal) folds and salivary ducts; G, Stomach opened with a mid-dorsal longitudinal cut
to expose inner anatomy.

VOLUME 94, NUMBER 2 609

a large cavity in the center of the foot. The cavity is lined with thick glan-
dular tissue embedded in the muscular mesopodium and is connected with
the pedal hemocoel. The organ and cavity taper toward the left side of the
foot. The ovipositor is innervated by a pair of nerves originating from the
metapodial ganglia (pg). Although the exact function of the ovipositor was
not observed, it is undoubtedly associated with the formation of the jelly
egg mass. In preserved specimens the cavity is filled with a viscous mucus
and continues to emit mucus when placed in water. There is no evidence
that it is a brood sac.

Nervous system (Fig. 4, 5B, C).—Diastoma has an epiathroid nervous
system that is moderately condensed with a layout typical of cerithiaceans.
The RPG ratio of Davis et al. (1976:263) (length of the pleurosupra—esoph-
ageal connective divided by the sum of the lengths of the supraesophageal
ganglion, pleuro-supraesophageal connective and right pleural ganglion) is
0.69, a median value between those observed in members of the Cerithiidae
and the Potamididae. The most characteristic features of the nervous system
are the long labial (Fig. 4, //n, din) and proboscis nerves which emerge from
the cerebral ganglia (rcg, cg). These lie free within the cephalic cavity in
a “‘loose’’ state and are surrounded with little connective or muscular tissue.
The large cerebral ganglia (Fig. 5B, C, cg) are almost fused to each other.
Each of the pleural ganglia (rpg, [pg) are joined to the cerebral ganglia with
short thick connectives. The right pleural ganglion (rpg) gives rise to a long
supraesophageal connective (sec) that ends in the supraesophageal ganglion
(spg) which is embedded _in the left body wall. This ganglion emits a nerve
that is connected to the left mantle nerve by a long dialyneury (d). The left
pleural ganglion (/pg) is joined to the subesophageal ganglion by a short
thick connective. A long visceral nerve (sbv) and a typical visceral loop are
present. The 2 pedal ganglia (pg) are joined to the cerebral and pleural by
moderately short, slender connectives. The pedal commissure is short and
thick. A large statocyst (st) containing numerous statoliths lies at the pos-
terior base of each pedal ganglion. Two long major connectives run from
each pedal ganglion to the propodial and mesopodial ganglia (Fig. 2C, mpg).
The mesopodial ganglia give rise to the nerves that innervate the ovipositor.
All ganglia are pink in color.

Systematics

Diastomidae Cossmann, 1894:322; 1906:173-174.—Wenz, 1940:749-750.—
Pchelintsev and Korobkov, 1960:159.—Franc (in Grassé) 1968:281.—
Griindel, 1976:71-75.—Ladd, 1972:27.—Wilson and Gillett, 1979:61.—
Rehder, 1980:33 (non Diastomidae Cossmann).

Diastomatidae (emend. pro Diastomidae) Ludbrook, 1971:31; 1978:112.

This family comprises many fossil species and one Recent one. It is char-
acterized by individuals having turreted, elongate shells with convex whorls

610 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

and ovate apertures with continuous outer lips and wide shallow anterior
canals. Former varices are normally present. A paucispiral operculum, tae-
nioglossate radula, paired salivary glands, stomach with style sac and gastric
shield, a deep mantle cavity with open pallial gonoducts, and the absence
of a penis are characteristic of the living animal.

Remarks.—The family, as originally proposed by Cossmann (1894:322-
323), included a group of fossil genera (Sandbergeria Bosquet, Cryptaulax
Tate, Exelissa Piette, Teliostoma Harris and Burrows, Aurelianella Coss-
mann) centered around the genus Diastoma Deshayes. Cossmann
(1906:173) subsequently treated the family in more detail and added the
genera (subgenera) Aneurychilus Cossmann and Cerithidium Monterosato.
With the exception of Cerithidium and Diastoma, all of these taxa represent
fossil groups. Cossmann appears to have been unaware that Diastoma is
still extant in southern Australia, represented by Diastoma melanioides
Reeve. This is probably due the the fact that Reeve (1849) originally placed
it in Mesalia, a turritellid genus.

The original concept of the family is somewhat heterogenous, exemplified
by Cossmann’s (1906:174) division of the family into ‘‘cerithid,”’ ‘‘rissoid”’
and “‘bacilliform’’ groups. The subsequent expansion of the family by the
inclusion of various other small-shelled groups has distorted the family con-
cept into an unnatural polyphyletic group. This is reflected in the literature
and has been recently acknowledged by Ludbrook (1971) and Grindel
(1976). A brief summary of the history of the family concept follows.

Initially, the family was expanded to include other genera of small-shelled
mesogastropods by Dall (1889:258), who, misled by Deshayes’ (1861:413)
statement of an affinity between Diastoma and the rissoids, also suggested
that Diastoma was related to Alaba and a Bittium species. It is likely that
Dall had never seen the fossil Diastoma species from the Paris Basin and
merely compared the figures of them with Alaba and the common south-
eastern Atlantic species, Bittium varium Pfeiffer. He was also unaware of
the existence of the living relict, Diastoma melanioides. Dall’s (1889) as-
signment of Bittium varium to Diastoma probably led other workers such
as Bartsch (1911), Wenz (1940) and Franc (1968) to equate cerithiacean
species of the genera Finella, Alaba, and Alabina, all having small shells
with shallow anterior canals which superficially resemble those of the large
fossil Diastoma species, with the Diastomatidae. Both Wenz (1940) and
Franc (1968) synonymized the Finellidae and the Alabinidae with the Dia-
stomatidae. Neither worker was aware that a living species of Diastoma
existed. This led more recent authors such as Ladd (1972:27), Kensley
(1973:281), and Abbott (1974:107) incorrectly to assign other small-shelled
cerithiacean snails of the genera Bittium, Alabina, Obtortio, and Diala to
the Diastomatidae.

VOLUME 94, NUMBER 2 611

Tate (1894:176—-177), Finlay and Marwich (1937:42), Marwick (1957:163),
and Ludbrook (1971) were the only authors to point out that Diastoma has
a living representative. Ludbrook (1971) also noted that many of the small
cerithiids referred to Diastoma should be excluded from that group. Her
treatment of the family, although brief, is substantially in agreement with
my findings.

Griindel (1976) suggested that the Diastomatidae was a primitive group
which arose in the Triassic and gave rise to the Cerithiacea. He believed
that the Cerithiidae split off from the Diastomatidae in the Cretaceous and
that Diastoma was a Short-lived unsuccessful side branch of mainstream
cerithiacean evolution. Within his classificatory scheme, Grtindel (1976:75)
divided the Diastomatidae into two subfamilies, the Diastominae, which
included the Bittiinae and Diastomiinae, and the Finellinae to which he
assigned the tribes Finellini and Scaliolini. Thus he included within the
subfamily Finellinae genera such as Sandbergeria, Eufinella, Obtortio,
Fesandella, and Scaliola. Grindel did not cite Ludbrook’s (1971) paper and
made no mention of the living species, Diastoma melanioides. His treatment
of the family placed much emphasis on sculptural characters of the proto-
conch and early teleoconch; moreover, he grouped genera solely on the
basis of morphological similarity. I thus find his conclusions unsatisfactory.
For instance, his assignment of Bittium to the Diastomatidae is incorrect
because anatomical evidence shows that Bittium is closely related to Cer-
ithium and falls within the Cerithiidae (see my recent remarks on this sub-
ject, Houbrick, 1980:4—5). The Finella group is also more properly assigned
to the Cerithiidae (Houbrick, 1980:4), and species in the Scaliola group
are anatomically unknown. Thus Griindel’s (1976) subfamily Fenellinae is a
polyphyletic group and should be excluded from the Diastomatidae.

The family name Diastomidae was correctly changed to Diastomatidae by
Ludbrook (1971:31) and the latter will be used throughout this paper. Abbott
(1974: 107), Griindel (1976:71), and Rehder (1980:33) all incorrectly dated the
family name from 1895, but Cossmann’s original proposal of the family was
published in 1894.

Grundel (1976:76), in a rather lengthy treatment of the Diastomatidae,
concluded that the genus Diastoma is closely related to Bittium, due to
similarity in shell features. His lack of acquaintance with the living species
and excessive reliance on ontogenetic characters derived from protoconch
and sculpture render his conclusions inadequate. Too much is known of the
plasticity of developmental modes of cerithiaceans to accept protoconch
shape and sculpture as conservative characters for phylogenetic specula-
tion. Grindel’s (1976:88) judgement that the presence of a well-defined an-
terior siphonal canal in Bittium species constitutes only a trivial difference
ignores the functional significance of such a difference. Bittium species are

612 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tiny snails that live on algae or hard substrata and represent a completely
different adaptive radiation from the Diastoma lineage, which is adapted
for burrowing in a sandy substrate.

Study of the living species, Diastoma melanioides, has supplied additional
characters derived from the radula and anatomy which more tightly define
the family limits. It is apparent that several of the small-shelled cerithi-
acean taxa discussed above should be excluded from the Diastomatidae and
that the family as defined herein, comprises an extinct Tethyan group with
a sole survivor living in south Australia. The exact status and placement of
Obtortio and Alaba remain unknown until more is known of their anatomy.

Diastoma Deshayes, 1850

Diastoma Deshayes, 1850:46; 1861:411-413 (Type-species, by monotypy,
Melania costellata Lamarck, 1804).—Tate, 1894:176—177.—Fischer,
1884:682—683 .—Harris, 1897:231—232.—Cossmann, 1889:34—35; 1906:175—
176.—Bartsch, 1911:581 (non Diastoma Deshayes).—Grant and Gale,
1931:758 (non Diastoma Deshayes).—Olsson, 1929:22 (non Diastoma
Deshayes).—Wenz, 1940:749-750 (in part).—Ludbrook, 1957:22 (n
part).—Pchelintsev and Korobkov, 1960:159.—Keen, 1963:32-86. (non
Diastoma Deshayes).—Abbott, 1974:107 (non Diastoma Deshayes).—
Emerson and Jacobson, 1976: (non Diastoma Deshayes).—Grundel,
1976:73-74, 76.—Houbrick, 1977:102; 1980a:3—4.—Ludbrook, 1978:112.

Neodiastoma Cotton, 1932:541 (Type-species, by original designation, Mes-
alia melanioides Reeve, 1849).—Finlay and Marwick, 1937:42.—Mar-
wick, 1957:162—163.—G6tting, 1974:129.

Description.—Shell turreted, elongate, of moderate size with inflated
whorls and ovate aperture that is broad at base and narrow posteriorly
where the outer apertural lip joins the body whorl to form a sutural notch.
Sculpture consists of axial ribs, thin spiral cords and occasional varices.
Suture deeply incised. Columella concave with slight median, oblique fold.
Anterior siphonal canal wide and shallow, not distinct from base of outer
apertural lip. Outer lip of aperture thin and slightly sinuous. Operculum
ovate, corneous, paucispiral, and with eccentric nucleus. Periostracum with
tiny hairs. Radula taenioglossate (2 + 1 + 1 + 1 + 2). Mantle cavity deep.
Ctenidium and osphradium do not extend length of mantle cavity. Salivary
glands tube-like and extending through the nerve ring. Style sac and gastric
shield present. Pallial gonoducts open in both sexes, males aphallate; sem-
inal receptacle and spermatophore pouch in outer lamina of pallial oviduct.
Females have a large ovipositor on right median part of mesopodium ex-
tending into center of foot and innervated by mesopodial ganglion. Nervous
system epiathroid, moderately condensed with long labial and proboscis
nerves and left dialyneury. Reproduction dioecious; spermatophores pres-
ent; spawn and larvae unknown.

VOLUME 94, NUMBER 2 613

Remarks.—This genus comprises a compact group of relatively large-
shelled, distinctive looking mesogastropods most of which are extinct. It is
now represented by a single living species in a very restricted area along
the coast of southern Australia (Fig. 6). Proposed by Deshayes in 1850, the
genus is best known from the Eocene of the Paris Basin. The original generic
citation in the Atlas of the Traité Elémentaire de Conchyliologie mentioned
only the name, Diastoma. The type-species is Melania costellata Lamarck,
1804, by monotypy. Deshayes (1861:411) later described the genus in con-
siderable detail from four Paris Basin species and discussed his reasons for
transferring Melania costellata to Diastoma. Deshayes (1861) further re-
marked that Diastoma only superficially resembled the melanians and that
the genus was closer to the Rissoidae. This has unfortunately led to con-
fusion about the generic limits and has resulted in a polyphyletic familial
concept, as discussed previously.

Tate (1894:176-177) was first to note that an Australian fossil, Diastoma
provisi Tate, 1894, was congeneric with the living Australian snail, Dia-
stoma melanioides. The latter was originally assigned by Reeve (1849) to
Mesalia Gray, a turritellid genus. Tate (1894) noted that it superficially
resembled Mesalia, but differed from other species in that genus by having
a columellar plait, a straight outer lip, and varices. He thus transferred
Mesalia melanioides to the genus Diastoma. Finlay and Marwick (1937:42)
assigned Diastoma to the Melantidae (=Thiaridae Troschel).

Cotton (1932:541) proposed the genus Neodiastoma which he assigned to
the Cerithiidae to accommodate the recent species, Diastoma melanioides.
He noted that this species differed from fossil Diastoma costellata by the
‘‘anterior notch of the outer lip’’ in the latter. Ludbrook (1971:31-—32) point-
ed out that not all specimens of Diastoma costellata have the aperture
separated at the suture to the same degree; thus, the presence of a sutural
ramp in association with the outer lip of the anterior aperture is a variable
feature. Moreover, examination of a series of Recent Diastoma melanioides
shows that this species also has a sutural ramp at the anterior aperture
although it is not as pronounced as in Diastoma costellata. | have figured
specimens of D. costellata with pronounced ramps (Fig. 1E—F). A single
variable shell character which is present in both fossil and Recent species
does not constitute a reasonable generic character. I concur with Ludbrook
(1971) that Neodiastoma is a synonym of Diastoma. Cossmann (1889:34)
originally suggested that the genus was similar to Semivertagus Cossmann
and Fastigiella Reeve in the placement of the median columellar fold, but
I think this is more easily explained by convergence. Cossmann (1906: 175)
later remarked that the genus was an “‘ambiguous’’ one and noted the su-
perficial similarity between Diastoma and the freshwater genus Melania.
He did not agree with Dall’s (1889) referral of Bittium varium Pfeiffer to
Diastoma and considered it a true Bittium species with a rudimentary canal
which superficially resembled that of Diastoma.

614 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fossil record.—According to Cossmann (1906:176), this genus may date
back to the Cretaceous (Senonian) but the assignment of fossils from this
Period to Diastoma, s.s., is unlikely. The genus, as defined herein, was
present in the Paleocene but is best known from the Eocene and Oligocene
of the Paris Basin. Diastoma species [Cossmann (1906) recorded 12] were
widespread in the Tethys Sea. Ludbrook (1971:31) recorded the genus from
Egypt, the East Indies, and North and South America, but gave no citations
to support this statement. Although I have made a literature search, I have
been unable to confirm the presence of Diastoma fossils in these areas. The
genus is found in the Australian Miocene, early Pliocene, and Pleistocene
where it is represented by three fossil species (Ludbrook 1971:31; 1978:112),
one of which, Diastoma melanioides, survives to the present.

Diastoma melanioides (Reeve, 1894)

Mesalia melanioides Reeve, 1849: Mesalia, fig. 3, sp. 3 (Holotype:
BM(NH); Type-locality, here selected: (Esperance, Western Australia).

Diastoma melanioides (Reeve), Ludbrook, 1971:32; 1978:112.

Diastoma melanoides Tate, 1894:177 (err. pro melanioides Reeve).

Mesalia exilis Sowerby, 1913:236, pl. 3, fig. 9 (Holotype: BM(NH); Type-
locality: ““West Australia.”’

Neodiastoma melanioides (Reeve), Cotton, 1832:541.

Synonymic remarks.—Reeve (1849) first described this species from an
unknown locality and remarked that “‘the generic pecularities of Mesalia
are well represented in this species.’’ This observation does not conform to
the facts: Mesalia species are anatomically unknown but are thought to be
members of the family Turritellidae and do not have the median columellar
fold seen in Diastoma melanioides. Sowerby later (1913:3, 236) described
the same species, naming it Mesalia exilis, but remarked that he was un-
certain of its generic allocation. He cited West Australia as the habitat but
Cotton (1932:541) noted that the type of Sowerby’s Mesalia exilis came
from Esperance Bay, Western Australia. Sowerby (1913) was either un-
aware of Reeve’s (1849) previous description of Mesalia melanioides or did
not regard Reeve’s species to be conspecific with Mesalia exilis. These two
taxa are, nevertheless, conspecific. Tate (1894) was the first to transfer
Reeve’s species from Mesalia to Diastoma and his decision was recon-
firmed by Cotton (1932:541) who elected to give the living form a different
generic name, Neodiastoma.

Ecology

Nothing has been recorded about the ecology of this species. Museum
data and observations made by Mr. A. Longbottom (pers. comm.) have
supplied the sparce information presented below. Diastoma melanioides
lives subtidally in depths of 1—5 meters where it burrows in clean white sand

VOLUME 94, NUMBER 2 615

ml
1200 KILOMETERS

Fig. 6. Geographic distribution of Diastoma melanioides based on museum specimens and
records cited by Ludbrook (1957). Star represents a fossil from the Ecula Basin.

associated with grass beds and algae. Its presence may be detected by long
trails in the sand. The large gastric shield and style sac indicate an herbiv-
orous mode of feeding; moreover, the tiny jaws, small taenioglossate radula,
stomach contents, and fecal pellets comprised of fine sand grains and de-
tritus suggest that Diastoma feeds on microalgae and detrital particles found
in the sandy substrate. Repaired shells indicated that this species is attacked
by crabs. Although no drilled shells were seen, the sandy habitat also sug-
gests possible predation by naticid snails. Paired snails were collected in
June (Longbottom, pers. comm.) and large ova were found in the ovaries
of preserved animals collected in July. This may indicate an early winter
breeding season. The spawn is undescribed, but the presence of a large
Ovipositor and associated mucoid-jelly gland suggests a jelly-like spawn

616 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

mass not unlike those described for other cerithiacean snails. The bulbous,
smooth protoconchs and restricted distribution are indicative of direct de-
velopment.

Ludbrook (1971:30) presented a map showing the summer isotherms of
the Great Australian Bight. These indicate that Diastoma lives well within
the warm temperature 19—20°C isotherms.

Conclusions

The shell and anatomy of Diastoma melanioides establish beyond doubt
that it is the superfamily Cerithiacea. Features common to the superfamily
are the open pallial gonoducts, aphallic males, and general layout of the
alimentary tract which includes a taenioglossate radula, paired salivary
glands and stomach with a style sac, gastric shield, and reduced spiral cae-
cum.

Distinctive features which establish the familial status of the group are
the elongate shell with its wide aperture, indistinct inhalant siphonal canal,
and deeply impressed suture and ramp. The shell shape is much like those
of the freshwater melanians. The thick, slightly hairy periostracum is an
unusual trait for a burrowing snail. I know of no other cerithiacean having
a similar periostracum although Campanile symbolicum, which is sympatric
with Diastoma, has a strange calcified periostracum (Houbrick, 1981a).

The anatomy of Diastoma differs markedly from other cerithiaceans by
a number of diagnostic features:

1) The highly extensible snout and large cephalic cavity are notable char-
acters. The snout of Diastoma is wider and longer in relation to body size
than in most cerithiaceans but there is some resemblance to burrowing,
sand-dwelling cerithiid snails such as Rhinoclavis fasciata (Bruguiere), R.
vertagus (Linnaeus) and R. aspera (Linnaeus) (Houbrick, 1978). The rela-
tively small buccal mass, jaws, and generalized taenioglossate radula are
not diagnostic at the familial level.

2) The paired worm-like salivary glands extend through the nerve ring,
are unusually long and simple, and constitute a distinctive character. In
contrast to Diastoma, the salivary glands of all observed cerithiid and mod-
ulid snails are spongy masses of tissue having long, thin ducts and, with the
exception of a small part of the left salivary gland, lie anterior to the nerve
ring (Houbrick, 1980:124). Some members of the Potamididae, such as Cer-
ithidea scalariformis (Say) and Batillaria minima (Gmelin) have more sim-
ple, worm-like salivary glands like Diastoma but the greater part of these
glands lies well anterior to the nerve ring (personal observation).

3) An unusual feature of the anterior and mid-esophagus of Diastoma is
its loose connection to the walls of the spacious cephalic cavity by numer-
ous, long muscular strands. The muscles are inserted on the esophagus

VOLUME 94, NUMBER 2 617

where it passes through the nerve ring and are also loosely connected to the
ganglia. The esophagus is thus able to move somewhat through the nerve
ring when the snout is extended. I have not seen this kind of anatomical
flexibility in the anterior esophagus of any other cerithiacean.

4) The pallial gonoduct of Diastoma has a simple, generalized arrange-
ment. The spermatophore pouch and seminal receptacle are located in the
outer lamina of the pallial oviduct and comprise a simple, bifurcate chamber.
Other cerithiaceans, such as the cerithiids, modulids, melanians, and tur-
ritellids, while having a similar ground plan to the pallial oviduct, display
greater complexity in the internal structures associated with the seminal
receptacle, spermatophore pouch and sperm collecting gutters.

5) An unusual diagnostic character of Diastoma is the large, complex
Ovipositor which is found on the mesopodium of females. Although ovipos-
itors have been noted in other marine cerithiacean snails such as Cerithium
(Marcus and Marcus, 1964:500) and Modulus (Houbrick, 1980b:121), I know
of no other marine snail in which there is a large mucus-producing gland
located within the center of the mesopodium and connected to the external
ovipositor. It is assumed that this gland contributes to the formation of the
egg mass, which is probably very gelatinous. Belgin (1973:390, fig. 7) has
shown a very similar ovipositor in Zemelanopsis, a freshwater melanian
snail, but gave no details about its internal structure other than to say that
it was a deep pit bordered by a muscular lobe.

6) A diagnostic feature of the nervous system of Diastoma is the length
of the labial and buccal nerves and the innervation of the ovipositor organ
by the mesopodial ganglion and its ancessory nerves. The nervous system
is moderately concentrated having an RPG ratio which lies between the
values observed in members of the Cerithiidae and Potamididae.

7) In Diastoma, the proximal ends of the osphradium, ctenidium, and
hypobranchial gland are not located at the posterior end of the mantle cav-
ity. Instead they begin anterior to it, which is an unusual feature. In mem-
bers of the Cerithiidae, Potamididae, Modulidae, and Turritellidae, these or-
gans extend the length of the mantle cavity, but in freshwater melanian
species of Zemelanopsis they appear to be shortened as in Diastoma (Bel-
gin, 1973:390, fig. 7).

While taxonomic criteria for the familial status of a group are ill-defined
and somewhat arbitrary, I believe that the characters derived from the shell
and anatomy of Diastoma are of sufficient diagnostic weight to establish the
group as a distinct family. To these I add the additional evidence of a long
fossil record which shows that these species comprised a distinct group
since the early Tertiary. It may be argued that the overall resemblance of
Diastoma to the cerithiids is quite close and that the evidence for familial
Status is not convincing. Nevertheless, the observed differences indicate to
me a higher taxonomic category for the group than a generic one.

618 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

The shell and anatomy of Diastoma are most like those observed in mem-
bers of the Cerithiidae and Potamididae and very much like those of some
species of the freshwater family Melanopsidae H. and A. Adams. It is pos-
sible that Diastoma is the marine branch of that family, but not enough
comparative material has been analyzed to say more on this subject. Al-
though this resemblance may be due to convergence, it is likely that the
Diastomatidae are closely related to these cerithiacean families. The shell
of Diastoma, in contrast to those of members of the above-mentioned fam-
ilies, lacks a distinct anterior canal and its aperture is more like those seen
in smaller-shelled members of the Dialidae. There may be a relationship
between these two groups but nothing is now known of the anatomy of
Diala.

The relationship between Diastoma melanioides and the Australian fossil
Diastoma species is clearly a close one (Ludbrook, 1971:131). There is no
way to demonstrate with certainty that this Australian assemblage belongs
to the same lineage as the Tethyan group of European fossils which bears
the same generic name. As is so often the case, the type-species of the
genus, Diastoma costellata, is rather atypical of the group. The axial ribs
are more pronounced and the sutural ramp, formed where the outer lip
meets the body whorl, is more exaggerated than in most Diastoma species
(see illustrations in Fig. 1E—F). Examination of a number of specimens of
this species shows that this character varies in intensity and is most pro-
nounced in older specimens. Moreover, other Tethyan Diastoma species
do not have this feature in so pronounced a state. It should also be noted
that a similar, but weaker sutural ramp is present in the living species and
was figured by Ludbrook (1971:fig. 13).

The main conchological difference between the Australian and European
fossils is the presence of the median columellar fold in species of the former.
This fold does not extend up the shell axis but appears to be merely a
thickening of the columellar callus. Cotton (1932), noting this difference
between the living species and the European fossils, considered it to be of
generic significance. I do not believe a simple character should be so heavily
weighted. Although the two assemblages may constitute different generic
taxa, I regard them as being of the same lineage and prefer to retain all
species with the genus Diastoma until more detailed analysis of the fossil
Species is made.

Members of the Diastomatidae probably lived throughout the Tethys Sea
during the Tertiary. Other Tethyan survivors are found in the Australian
region and include prosobranchs such as Campanile symbolicum Iredale
(Houbrick, 1981a), Gourmya gourmyi (Crosse) (Houbrick, 1981b), and the
bivalve genus Neotrigonia, family Trigoniidae, which became largely extinct
in the Cretaceous. These relict species probably remained in the Australian
area after the closure of the Tethys in the Miocene.

VOLUME 94, NUMBER 2 619

Acknowledgments

I am indebted to Mr. Alan Longbottom of Esperance, Western Australia,
for sending me the preserved animals upon which this paper is based. I
thank Dr. Fred Wells of the Western Australian Museum, Perth, for his
assistance in this project and for the study of specimens in his charge. Dr.
Winston Ponder of the Australian Museum, Sydney, kindly allowed me to
study Diastoma specimens in that collection. I am grateful to him for his
critical reading of a draft of this paper. I also thank Drs. Harald Rehder and
Joseph Rosewater of the National Museum of Natural History, Smithsonian
Institution, and Dr. Kenneth J. Boss of the Museum of Comparative Zo-
ology, Harvard University, for their comments and suggestions. Thanks are
extended to the staff of the Scanning Electron Microscope Laboratory and
to Victor Krantz of Photographic Services, Smithsonian Institution, for the
photographs.

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Bartsch, P. 1911. The Recent and fossil mollusks of the genus Diastoma from the west coast

of America.—Proceedings of the United States National Museum 39:581—584.

. 1911. The Recent and Fossil Mollusks of the Genus Alabina from the west coast of

America.—Proceedings of the United States National Museum 39:409-418, pls. 61-62.

Belgin, F. H. 1973. Studies on the functional anatomy of Melanopsis praemorsa (L.) and
Zemelanopsis trifasciata (Gray). Proceedings of the Malacological Society of London
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Cossmann, M. 1889. Catalogue illustré des Coquilles Fossiles de l’;Eocéne des environs de

Paris.—Annales de la Société Royale Malacologique de Belgique 24:3-381, pls. 1-9.

1894. Revision sommaire de la faune du Terrain oligocene marin aux environs

d’ Etampes.—Journal de Conchyliologie 41:297-363.

. 1906. Essais de Paleoconchologie Comparee 7:261. Paris.

Cotton, B. C. 1932. Notes on Australia Mollusca with descriptions of new genera and new
species.—Records of the South Australian Museum 4(4):537—-547.

Dall, W. H. 1889. Reports on the results of dredging, under the supervision of Alexander
Agassiz, in the Gulf of Mexico (1877-78) and in the Caribbean Sea (1879-80), by the U.
S. Coast Survey Steamer ‘“Blake,’’ Lieut.-Commander C. D. Sigsbee, USN, and Com-
mander J. R. Bartlett, USN, commanding. Report on the Mollusca. Part 2 Gastropods
and Scaphopoda.—Bulletin of the Museum of Comparative Zoology 18(29):1—492, 40
pls.

Deshayes, G. P. 1850. Traité Elémentaire de Conchyliologie, Atlas, 80 pp., 132 pls.

. 1861-1864. Description des Animaux sans Vertebrés découverts dans la Bassin de

Paris. Paris. vol. 2, text: Mollusques, Acéphales monomyaires et Branchiopodes.—Mol-

lusques Céphalés, premiere partie, 968 pp., atlas, 64 pls.

Emerson, W. K., and M. K. Jacobson. 1976. The American Museum of Natural History
Guide to Shells: Land, Freshwater, and Marine, from Nova Scotia to Florida.—New
York. i—xviii + 482 pp., 17 pls.

Finlay, H. J., and J. Marwick. 1937. The Wangaloan and associated Molluscan Faunas of
Kaitangata—Green Island Subdivision.—New Zealand Geological Survey Branch. Pa-
laeontological Bulletin No. 15:1140, 17 pls.

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Franc, A. 1968. Classe des Gastéropodes (Gastropoda Cuvier, 1798) [in] Grassé, P. P. (ed.),
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Fretter, V., and A. Graham. 1962. British prosobranch molluscs: their functional anatomy
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Gotting, K. 1974. Malakozoologie, Grundriss der Weichtierkunde.—Stuttgart, 320 pp.

Grant, U. S., and H. R. Gale. 1931. Catalogue of the Marine Pliocene and Pleistocene Mol-
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Griindel, J. 1976. Zur Taxonomie und Phylogenie der Bittium-Gruppe (Gastropoda, Cerithi-
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Harris, G. F. 1897. Catalogue of Tertiary Mollusca in the Department of Geology, British
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Houbrick, R. 1977. Reevaluation and new description of the genus Bittium (Cerithiidae).—

The Veliger 20(2): 101-106.

1978. The family Cerithiidae in the Indo-Pacific, Part I: The genera Rhinoclavis,
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. 1980b. Observations on the anatomy and life history of Modulus modulus (Proso-

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198la. Anatomy, biology and systematics of Campanile symbolicum Iredale with
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. 1981b. Anatomy and Systematics of Gourmya gourmyi (Prosobranchia: Cerithiidae),

a Tethyan Relict from the south west Pacific.—The Nautilus 95(1):2-11.

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of Western Australia.—Bulletin 125:1-286, 24 pls.

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Marcus, E., and E. Marcus. 1964. On Cerithium atratum (Born, 1778).—Bulletin of Marine
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VOLUME 94, NUMBER 2 621

Pchlintsev, V. F., and I. A. Korobkov. 1960. Molliuski-Briukhonogie. In Y. A. Orlov (ed).—
Osnovy Paleontologii, Moskva, 359 pp.

Reeve, L.A. 1849. Conchologia Iconica, 5, Mesalia, 1 pg, 1 pl.

Rehder, H. A. 1980. The Marine Mollusks of Easter Island (Isla de Pascua) and Sala y
Gomez.—Smithsonian Contributions to Zoology No. 289:1-—167, 14 pls.

Sowerby, G. B. 1913. Descriptions of New Species of Mollusca.—Annals and Magazine of
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Tate, R. 1894. Unrecorded genera of the older Tertiary Fauna of Australia, including diagnoses
of some new genera and species.—Journal of the Royal Society of New South Wales
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dewolf, Handbuch der Palaozoologie, vol. 6.—Borntraeger, Berlin. viit+ 1639 pp.
Wilson, B.,and K. Gillett. 1979. A Field Guide to Australian Shells.—Prosobranch Gastropods.

287 pp., 67 pls. Sydney.

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

PROC. BIOL. SOC. WASH.
94(2), 1981, pp. 622-630

PSEUDOPLUMARELLA ECHIDNA, A NEW SPECIES
OF PRIMNOID OCTOCORAL FROM QUEENSLAND
(COELENTERATA: OCTOCORALLIA)

Frederick M. Bayer

Abstract.—A new primnoid octocoral, Pseudoplumarella echidna, from
the southern coast of Queensland, is described and illustrated by scanning
electron micrography. The species is compared with the related Pseudoplu-
marella corruscans (Thomson and Mackinnon, 1911) from New South
Wales, which also is illustrated. A possibly distinct variant is reported but
not established as a distinct taxon.

A brief examination of octocorals in the collections of the Queensland
Museum, made possible through the kindness of Dr. L. R. G. Cannon,
revealed an interesting new species of the primnoid genus Pseudoplumarella
allied to P. corruscans (Thomson and Mackinnon, 1911). As the octocoral
fauna of Australia is still inadequately known, it is desirable to put on record
descriptions of new species such as this in order to facilitate a comprehen-
sive marine faunal inventory for Australia at some time in the future.

Pseudoplumarella Kukenthal, 1915

Plumarella (part) Thomson and Mackinnon, 1911:682.
Pseudoplumarella (part) Kukenthal, 1915:143, 145; 1924:263.

Diagnosis (emend.).—Primnoids with uniplanar, alternately pinnate
branching. Polyps small, directed obliquely upward, biserial or on all sides
of twigs; lateral and adaxial rows of body scales greatly reduced, adaxials
absent in some species; opercular scales tall isosceles triangles, of nearly
uniform size, only the adaxials being somewhat smaller, fitting closely to-
gether without overlapping, forming a prominent conical operculum; mar-
ginal scales fewer than 8, not folding over operculars; coenenchymal scler-
ites scalelike, sometimes with a deeper layer of small, tuberculate spheroids.

Type-species.—Plumarella thetis Thomson and Mackinnon, 1911; by orig-
inal designation.

Remarks.—Although Kukenthal (1915, 1924) included Plumarella plu-
matilis (Milne Edwards and Haime, 1857) in his genus Pseudoplumarella,
Versluys (1906:39) had already established the subgenus Pterostenella for
that species. Therefore, were it not for the fact that P. plumatilis and P.

VOLUME 94, NUMBER 2

Fig. 1. Pseudoplumarella echidna, colonies: A, Holotype, Queensland Mus. Reg. No.
G4710; B, Paratype, Queensland Mus. Reg. No. GL1; C, Paratype, Queensland Mus. Reg. No.
GL2; D, Paratype, USNM 59823; E, Variant, Queensland Mus. Reg. No. GL3.

624 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. A, Pseudoplumarella echidna, holotype; B, P. echidna, paratype, GL1; C, P.
echidna, variant GL3; D, P. corruscans, syntype, BM(NH) 1933.3.13.73. Parts of branches
with polyps; all SEM micrographs.

thetis are generically distinct, Pseudoplumarella would be a junior syn-
onym. The most conspicuous distinguishing character of Pseudoplumarella
is the tall, conical operculum composed of tightly fitting triangular scales
the margins of which abut but do not overlap; the polyps may be in close
spirals all around the twigs, or biserial, but not in whorls as in Pterostenella.
The operculum of Pterostenella is very low, often almost flat, and is com-
posed of overlapping scales; the polyps, which have a complete adaxial
covering of scales, stand almost vertically and are arranged in widely spaced
whorls.

In addition to the new species here described, the genus Pseudopluma-

VOLUME 94, NUMBER 2 625

Fig. 3. Pseudoplumarella echidna: Lateral and abaxial views of polyp; SEM stereomicro-
graphs, 50x.

relia includes four species: P. thetis, P. corruscans, P. filicoides, and P.
versluysi, all of Thomson and Mackinnon (1911) and all from the coast of
New South Wales, Australia. Unfortunately, ambiguities in the original de-
scriptions and discrepancies between descriptions and illustrations preclude
the construction of a reliable key for their separation. However, the new
species is easily distinguished from the others as it is the only one that has
spines on the abaxial marginal scales.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4 Pseudoplumarella echidna: A, Lateral view of intact polyp; B, Marginal scales, inner
and outer surfaces; C, Opercular scales; D, Adaxial opercular scale; E, Body scales; F, Sub-
marginal scale, outer surface; G, Submarginal scale, inner surface; H, Coenenchymal scales.

A, 38x; B-H, 63x.

Pseudoplumarella echidna, new species
Figs. la—d; 2a—b; 3-4

Material.—Four colonies, dredged amongst fan corals, Jumpin Pin, be-
tween North Stradbroke and South Stradbroke Islands, Queensland, where

VOLUME 94, NUMBER 2 627

Fig. 5. Pseudoplumarella corruscans, syntype, BM(NH) 1933.3.13.73; lateral and abaxial
views of polyp; SEM stereomicrographs, 50x.

southern Moreton Bay enters the ocean; 47 fathoms (86 m); coll. Prof.
William Stephenson and student party, | July 1961.

Diagnosis.—Pseudoplumarella with distal edge of marginal scales pro-
duced into strong spine.

Description.—Colonies (Fig. la—d) closely and alternately pinnate, 18-21
twigs in 5 cm along one side of branch; fully developed twigs generally of
rather uniform length throughout colony or within major branches, from 1.5
to 3 cm long, decreasing in length abruptly toward tips of branches. Polyps
small, about 0.75 mm tall, closely biserial on unbranched twigs; operculum
of one polyp often overlapping base of next polyp above; 17-20 in 1 cm
along one side of twigs (Fig. 2a). Operculum tall, conical, composed of
triangular scales closely fitting but not overlapping (Fig. 3), size nearly uni-
form (Fig. 4c) except for adaxial pair, which are somewhat smaller than the
others (Fig. 4d). Marginal scales usually 4, in abaxial and outer lateral rows
only, rarely a small scale below one or both inner lateral operculars; free
edge of marginal scales produced as a sharp spine, that of abaxial marginals

628 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Pseudoplumarella echidna, variant; lateral and abaxial views of polyp; SEM ste-
reomicrographs, 50x.

stronger (Figs. 3, 4b) that of the outer laterals shorter and usually not so
acute. Usually 5 (sometimes 4, rarely 6 or 3) body scales in abaxial rows,
usually only 1 or 2 in outer lateral rows; body scales (Fig. 4e) oval; distal
edge of submarginals sometimes produced as a low, blunt spine or rounded
angle (Fig. 4f); body scales externally smooth except for some low, simple
granulations on proximal part, internally tuberculated, distal edge with sev-
eral sharp radial ridges (Fig. 4g). Tentacles devoid of sclerites. Coenenchy-

VOLUME 94, NUMBER 2 629

Table 1.—Comparison of measurements of Pseudoplumarella curruscans and P. echidna.

No. of Length Length No. of body
Height of twigs of twigs Polyps of polyp scales in
colony (cm) in 5 cm (cm) in 1 cm (mm) abax. rows
P. corruscans 44 13 4 10 1 6
P. echidna G4710 24.5 18 3 17 0.75 3-6
P. echidna GL1 22 20 S25)
P. echidna GL2 14.5* 21 2.5-3.0 20 0.75 3-4
P. echidna 59823 11.5 21 2 20 0.75 3-6
P. echidna var. Soe 31 ee) 14 0.75 3-4

* Broken off at top and lacking holdfast.

mal sclerites (Fig. 4h) scalelike, rounded, squarish, broadly or narrowly
oval, imbricate, their free edges upturned and outer face more or less con-
cave, thus resembling shallow, lopsided bowls with tuberculate bottoms.
Tuberculate spheroids were not observed.

Types. —Queensland Museum Register No. G4710 (holotype, Fig. 1a);
GL1I (paratype, Fig. 1b); and GL2 (paratype, Fig. 1c). USNM 59823 (para-
type, Fig. 1d).

Variant.—Queensland Museum Register No. GL3 (Fig. le).

Comparisons.—In gross colonial aspect, Pseudoplumarella echidna most
closely resembles P. corruscans (Thomson and Mackinnon), but in that
species the colonies are larger and the lateral twigs not so closely crowded.
Although Kiukenthal (1924:263) separated P. corruscans from P. thetis be-
cause ““Die abaxialen Schuppen sind schmaler und haben einen freien ge-
zahnelten Rand,’’ this character must have been based on Thomson and
Mackinnon’s pl. 65, fig. 4, which does not agree with the syntype (Fig. 5)
deposited in the British Museum (Nat. Hist.). Moreover, the original de-
scription makes no mention of a toothed margin on the body scales, nor
does the drawing of sclerites (pl. 68, fig. 6) illustrate such a feature. Possibly,
the “‘strongly-marked radiating ridges’’ on the inside border of the body
scales (which are visible in the intact polyps as can be seen in Fig. 5) led
to the erroneous drawing of the polyps.

Pseudoplumarella echidna also resembles the Japanese Plumarella spi-
nosa Kinoshita, 1908. That species has similar, closely pinnate branching
and small, biserial polyps with spinous marginal scales, but there are 8
marginals, 6 of them with spines, and the operculars overlap in such a way
that the operculum forms only a low cone or is nearly flat.

The polyps of P. corruscans are mostly about 1 mm tall (Fig. 5) and they
do not overlap one another along the branch (Fig. 2d), so about 10 occur in
1 cm along one side of a branch, compared with 17—20 in P. echidna.

The measurements of Pseudoplumarella corruscans as presented by
Thomson and Mackinnon (1911:684) are compared with those of the present
specimens in Table 1.

630 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

A single specimen from the haul that yielded the holotype and paratypes
of P. echidna differs in features that may be of specific importance, but the
lack of additional specimens precludes establishing it as a new species at
this time. It differs from P. echidna in the following ways: the colony (Fig.
le) is smaller (a little over 5 cm tall, but in damaged condition), the twigs
shorter (1.5 cm) and more crowded (31 in 5 cm along one side of a branch),
and the polyps less crowded (14 in | cm along one side of twig) (Fig. 2c);
in many polyps the abaxial body scales are reduced to a single row of 3 or
4, the marginal spines are not so strong, and the coenenchymal scales fit
tightly, with little or no overlap (Fig. 6). Differences of this magnitude would
ordinarily be considered of specific importance. However, the size differ-
ences could be related to the immediate environment of that colony, and
the reduction of the abaxial body scales to a single row could be an indi-
vidual anomaly, so this specimen is treated as a possible variant of Pseu-
doplumarella echidna pending the study of adequate material.

Acknowledgments

I am grateful to Dr. L. R. G. Cannon of the Queensland Museum, who
made these specimens available for study, and to the Queen Elizabeth Fel-
lowship Program, and especially to Drs. J. T. Baker, John Pollard, and
Shirley Dryden, who made it possible for me to carry out a highly productive
research program in Australia during June—September 1980.

The scanning electron micrographs reproduced herein were made by Wal-
ter R. Brown, Chief of the S.E.M. Laboratory, Smithsonian Institution, and
the photographs of colonies were made by Michael R. Carpenter, who also
assisted in the final preparation of the plates.

This paper has benefitted from the criticisms of my colleagues Drs. W.
Duane Hope and Raymond B. Manning, to whom I am grateful for time
spent in my behalf.

Literature Cited

Kinoshita, K. 1908. Primnoidae von Japan.—Journal of the College of Science, Imperial
University of Tokyo 23(12):1-74, pls. 1-6.

Kitkenthal, W. 1915. System und Stammesgeschichte der Primnoidae.—Zoologischer An-

zeiger 46(5):142—158.

. 1924. Gorgonaria.—Das Tierreich 47:i-xxviii + 1-478, 209 figs. Berlin and Leipzig,

Walter de Gruyter & Co.

Thomson, J., and D. L. Mackinnon 1911. The alcyonarians of the ‘‘Thetis’’ Expedition.—
Australian Museum, Sydney, Memoir 4:661—695, pls. 41-82.

Versluys, J. 1906. Die Gorgoniden der Siboga Expedition II. Die Primnoidae.—Siboga-Ex-
peditie Monographie 13a:1—187, figs. 1-178, pls. 1-10, chart.

Department of Invertebrate Zoology, National Museum of Natural His-
tory, Smithsonian Institution, Washington, D.C. 20560.

PROC. BIOL.-SOC. WASH.
94(2), 1981, pp. 631-639

SOLENOCERA ALFONSO, A NEW SPECIES OF
SHRIMP (PENAEOIDEA: SOLENOCERIDAE)
FROM THE PHILIPPINES

Isabel Pérez Farfante

Abstract.—A new species of penaeoid shrimp, Solenocera alfonso, is
described and illustrated from specimens collected in the waters of the Phil-
ippines at depths between 176 and 547 m. This species possesses a spine on
the dorsal midline of the carapace posterior to the cervical sulcus, a feature
which distinguishes it from all of its congeners. It is compared with the
Indo-West Pacific S. australiana, S. halli, and S. melantho.

Among the interesting materials encountered during a current study of
the extensive collections of penaeoid shrimps obtained in the Philippines by
the U.S. steamer Albatross, 1907-10, were representatives of a previously
unknown species of Solenocera. This shrimp, which is now known to occur
on the upper slope of the island platforms to depths of 547 m, is described
and illustrated here. It is characterized by the presence of a spine on the
dorsal midline of the carapace posterior to the level of the dorsal end of
the cervical sulcus, a feature unique within the genus. This character, to-
gether with the hepatic and branchiocardiac sulci being curved in opposing
arcs and the peculiar terminal armature of the ventromedian lobule of the
petasma, make this species stand out from its relatives. In recent years,
large collections of penaeoids have been assembled from Indonesia and the
Philippines, and I hope that specimens of this shrimp will be discovered
among these collections, thus expanding our knowledge of its geographical
and ecological distribution.

Type-material is deposited in the National Museum of Natural History,
Smithsonian Institution (USNM), and the Ryksmuseum van Natuurlijke
Historie (RMNH), Leiden. The method of measuring the specimens and the
terminology used below are described by Pérez Farfante (1969), and Pérez
Farfante and Bullis (1973). In the captions of the figures, carapace length
and total length are abbreviated cl and tl, respectively. The illustrations of
the petasma were made from a specimen stained with methyl green.

Solenocera alfonso, new species
Figs. 1-5

Material (all from the Philippines)—Holotype: 2, USNM 184022, 31.5
mm carapace length, about 114 mm total length, 10 mm rostrum length;

632 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Solenocera alfonso, holotype, USNM 184022, 2 31.5 mm cl, off Capitancillo I, W
of Leyte, Philippines: Lateral view. Scale = 10 mm.

type-locality, off Capitancillo I, W of Leyte, 10°38’00"N, 124°13'08"E, 346
m, 18 March 1909, Albatross stn 5409. Allotype: 6, USNM 184023, 26 mm
carapace length, about 109 mm total length, 8.5 mm rostrum length, Ma-
cajalar Bay, N Mindanao, 8°37'37"N, 124°35'00’E, 391 m, 4 August 1909,
Albatross stn 5501.

Paratypes: | 2, USNM 184024, Balayan Bay, Luzon, 329 m, 20 February
1909, Albatross stn 5363. 1 6, USNM 184025, Tayabas Bay, SE Luzon,
357 m, 24 April 1908, Albatross stn 5222. 1 2, USNM 184026, SE of San
Andrés Is, NW of Marinduque, 91-353 m, 24 April 1908, Albatross stn 5221.
1 2, USNM 184027, NE of Cebu, 130-271 m, 3 April 1908, Albatross stn
5194.2 6 1 9, USNM 184028, off Palompon, W Leyte, 344 m, 16 March
1909, Albatross stn 5402. 2 d6 1 29, USNM 184029, off Palompon, W of
Leyte, 333 m, 16 March 1909, Albatross stn 5403. 1 6 1 2, RMNH, N of
Ponson I, W of Leyte, 348 m, 17 March 1909, Albatross stn 5404. 1 9,
USNM 184030, N of Capitancillo I, W of Leyte, 291 m, 18 March 1909,
Albatross stn 5408. 1 6, USNM 184031, S of Panay I, 176 m, 30 March
1908, Albatross stn 5183. 1 6 2 2, USNM 184032, NE of Pescador I, E of
Negros, 547 m, 1 April 1908, Albatross stn 5189.3 ¢ 1 2, USNM 184033,
NE of Pescador I, E of Negros, 549 m, 1 April 1908, Albatross stn 5188. 1

VOLUME 94, NUMBER 2 633

Fig. 2. Solenocera alfonso, paratype, USNM 184026, 2 40 mm cl, SE of San Andrés Is,
NW of Marinduque, Philippines: Terminal parts of right antennular flagella—A, Dorsal: B,
Ventral. Scale = 2 mm.

6 1 2, USNM 184034, S of Dumalag I, Davao Gulf, 247 m, 18 May 1909,
Albatross stn 5247.

Description.—Body glabrous (Fig. 1) except for elongate patch of setae,
typical of members of the genus, covering rostrum immediately dorsal to
adrostral carina and continuing posteriorly to epigastric tooth. Rostrum al-
most reaching distomesial margin of eye, horizontal or tilted upward, deep,
ventral margin convex along basal 0.7—0.8, apically almost straight. Number
of rostral plus epigastric teeth 6-8 (usually 7), teeth progressively smaller
and closer together from epigastric to ultimate, latter distinctly postapical;
apex of fourth tooth in line with orbital margin. Adrostral carina, extending
from orbital margin to or slightly beyond ultimate tooth. Postrostral carina
well marked but comparatively low, and weakly depressed at level of dorsal
extremity of cervical sulcus, bearing tooth slightly anterior to or at mid-
length between depression and midposterior margin of carapace; carina pro-
gressively lower behind tooth, and followed by minute dorsal tubercle lo-
cated on depressed posteromedian portion of carapace. Median sulcus
varying from well marked and almost continuous to being represented only
by few pits. Marginal ridge of carapace narrow anteriorly, broadening and
forming ventral lobe at about midlength of branchial region, and broadening
again forming posterolateral lobe, finally becoming slender along midpos-
terior section. Orbital spine small; postorbital spine longest of lateral cara-

634 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Solenocera alfonso, allotype, USNM 184023, 5 26 mm cl, Macajalar Bay, N Min-
danao, Philippines: A, Dorsal view of petasma; B, Ventral view of same. Scale = 2 mm.

pace spines, strong and continuous with conspicuous buttress; antennal spine
slender; hepatic spine sharp. Pterygostomian and branchiostegal spines lack-
ing. Orbito-antennal sulcus weakly defined. Cervical carina raised, sharp, with -
distinct bend just dorsal to hepatic region, its posterodorsal extremity ending
at lateral base of epigastric tooth; cervical sulcus deep, merging ventrally
with anterodorsally directed shallow groove and hepatic sulcus, junction
forming deep depression just anterior to hepatic spine. Hepatic sulcus with
posterior part, relatively shallow but well defined, turning ventrally in arc;
anterior part deep and extending anteroventrally to pterygostomian pit; he-
patic carina sharp anteriorly, its anteroventral extremity making sharp bend
ventrally extending along posterior margin of tear shaped pterygostomian
pit. Branchiocardiac sulcus and carina strongly defined, forming broad an-
terior arc and extending posterodorsally to near margin of carapace.
Antennular peduncle about 0.6 as long as carapace; prosartema reaching
or almost reaching distomesial margin of eye, but its long setae attaining
proximal third of second antennular article. Stylocerite relatively short,
length equivalent to 0.65 that of first antennular article, and ending in in-
conspicuous spine; distolateral spine slender and sharp. Dorsal flagellum
slightly longer than ventral, about 1.6 times carapace length while ventral
flagellum 1.4 times carapace length in shrimp 23 mm cl; in shrimp 37 mm cl

VOLUME 94, NUMBER 2 635

dorsal 0.9 and ventral 0.7 times carapace length; terminal part of dorsal
flagellum gradually tapering distally over 9 to 24 (latter in young) articles
(Fig. 2A), that of ventral flagellum over 8 to 12 (Fig. 2B), in both flagella
last few articles forming filament.

Scaphocerite overreaching antennular peduncle by 0.1 to 0.2 of its own
length; lateral rib ending in strong, sharp spine, reaching or almost reaching
distal margin of lamella. In available specimens, antennal flagellum as much
as 4.1 times total length of shrimp.

Third maxilliped surpassing antennular peduncle by length of dactyl and
about half that of propodus; ratio of dactyl/propodus 0.8-0.9.

First pereopod extending to distal end of carpocerite or exceeding it by
as much as 0.8 length of propodus. Second pereopod overreaching anten-
nular peduncle by at least 0.75 length of propodus and at most by propodus
and extreme distal part of carpus. Third pereopod surpassing antennular
peduncle by length of propodus and 0.3 to 0.5 that of carpus. Fourth per-
eopod overreaching peduncle by at least half length dactyl and at most by
dactyl and 0.1 length of propodus. Fifth pereopod overreaching antennular
peduncle by length of dactyl and almost entire propodus or by dactyl, prop-
odus, and extreme distal part of carpus. Order of pereopods in terms of
their maximum anterior extensions: first (shortest), fourth, second, fifth,
and third—latter exceeding fifth only slightly if at all. Third maxilliped barely
Overreaching second pereopod. First pereopod armed with long, slender
mesial spine on basis and slightly shorter one on ischium. Second pereopod
bearing short spine on basis. In male, fifth pereopod bearing flattened, scale-
like tooth on distomesial margin of coxa; in female short tooth borne on
distomesial extremity of coxal plate.

Abdomen with middorsal carina from second through sixth somites, ca-
rina on second somite low and rounded (sometimes almost indistinct), that
on third to sixth, high and sharp. Posterodorsal margin of third through fifth
somites with median incision; sixth somite bearing conspicuous sharp spine
at posterior end of carina and pair of small posteroventral spines. Telson
tapering caudally to acute apex and bearing pair of small, fixed, lateral
spines. Mesial ramus of uropod approaching apex of telson; lateral ramus
reaching or barely surpassing distal border of mesial ramus, its marginal
ridge ending in well developed spine. All sternites armed with laterally com-
pressed, strong median tooth on posterior border, teeth decreasing in size
posteriorly.

Petasma (Fig. 3A—B; Fig. 4B) with distal part of ventromedian lobule
tapering rapidly and extending conspicuously beyond dorsolateral lobule,
latter surpassing distally and overlapping ventral costa dorsally. Terminal
margin of dorsomedian lobule unarmed, that of ventromedian lobule sinuous
and bearing 10 to 18 irregular spinules of variable length. Distal part of
dorsolateral lobule with lateral margin weakly biconcave, its terminal margin

636 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Solenocera alfonso, allotype: A, Dorsolateral view of right appendix masculina; B,
Dorsolateral view of distal part of left half of petasma. Scales = 1 mm.

armed with | or 2 well developed spinules at mesial extremity, and row of
very minute ones (usually lost) extending onto lateral margin; lobule with
stiff ridgelike thickening running obliquely from distal to proximolateral
margins; thickening becoming obliquely truncate proximally, abutting dis-
tomesial border of proximal part of ventral costa. Ventral costa bent strong-
ly outwardly, resulting flap, broadest mesially, and bearing 15 to 20 short
spines on terminal margin. Junction of distal and proximal parts of ventral
costa marked by deep emargination.

Appendix masculina (Fig. 4A) broadening from base to about midlength
then tapering gradually to blunt apex, and bearing dorsolateral, strongly
curved rib proximally; distal part of lateral margin armed with elongate
patch of long setae, and apex with tuft of shorter ones. Appendix interna
considerably narrower but extending as far as or slightly farther than ap-
pendix masculina, bearing apically closely set short setae. Basal sclerite
with subvertical lateral wall projecting distally in blunt ventrolateral spur.

Thelycum (Fig. 5) with subquadrangular posterior depression delimited
laterally by raised margins of sternite XIV and anteriorly by transverse
ridge, sometimes interrupted, situated somewhat anterior to midlength of
sternite; pair of slender, fusiform, submesial protuberances placed relatively
far apart, immediately anterior and subperpendicular to transverse ridge
(protuberances almost indistinguishable in young). Sternite XIII with high

VOLUME 94, NUMBER 2 637

Fig. 5. Solenocera alfonso, holotype: Thelycum. Scale = 2 mm.

vertical posterior shelf divided by median slit and bearing pair of strong,
acute, anterolateral tubercles; base of anterior third of sternite XIII crossed
by transverse elevation continuous with caudomedian ridge. Posterior tho-
racic ridge with thick, setose, strongly biconvex anterior border.

Size.—Carapace lengths of males examined 15 to 29 mm, of females, 18
to 40 mm.

Geographic and bathymetric ranges.—This species has been found only
in the waters of the Philippines: from Balayan Bay, SW Luzon, to Davao
Gulf, SE Mindanao. It inhabits the upper slope of the island shelves, at
depths of 176 to 547 m, and occurs on substrates of mud—mostly green
mud—and at least occasionally on a mixture of fine sand and mud on which
one of the 14 samples available was collected.

Remarks.—Solenocera alfonso is unique among the members of the genus
in possessing a spine on the postrostral carina posterior to the cervical
sulcus, about midway between the latter and the caudal margin of the car-
apace. In this species both the posterior part of the hepatic sulcus and the

638 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

anterior part of the branchiocardiac sulcus turn ventrally forming opposing
arcs which are neither so well defined nor present together in any of its
congeners. Also distinctive is the marginal carina of the carapace, which is
expanded in two lobes on each side: one situated posteroventrally
about midlength of the branchial region, the other posterolaterally near mid-
dorsum.

This species closely resembles members of a homogeneous group of Indo-
west-Pacific species comprising S. australiana Pérez Farfante and Grey,
1980, S. halli Starobogatov, 1972, and S. melantho De Man, 1907. This
group, like S. alfonso, may be characterized by lacking branchiostegal and
pterygostomian spines, possessing a hepatic carina ending at the pterygo-
stomian region (not recurving posteriorly), and having a relatively low post-
rostral carina that is only slightly depressed at the level of the cervical
sulcus. Solenocera alfonso may be distinguished from the other three
species not only by the peculiar features cited in the previous paragraph,
but also by thelycal and petasmal characters.

In S. alfonso the distal part of the ventromedian lobule of the petasma
tapers rapidly instead of gradually or being truncate, its free margin is sin-
uous and bears spinules of various lengths the number of which ranges from
10 to 18. In S. australiana, S. halli, and S. melantho the free margin of the
ventromedian lobule is armed with numerous spinules which in the former
two species increase in length laterally and in S. melantho are all about the
same length, except for the lateralmost three or four that may be smaller.
In S. alfonso the distal margin of the dorsolateral lobule at its mesial ex-
tremity bears only | or 2 readily distinguishable spinules continuous with
a row of very minute ones which extend along the lateral margin (these
Spinules are missing in most of the specimens available, there remaining
only the raised points where they were inserted). In S. australiana and S.
halli the dorsolateral lobule bears numerous spinules (18 to 40) which de-
crease in length proximolaterally, and in §. melantho, although the margin
may be armed with only | or 2 spinules at its mesial extremity—much like
in S. alfonso—there are usually more, as many as 13; furthermore, the
lobule itself is not stiffened by a ridgelike thickening as it is in S. alfonso.

The thelycum of S. alfonso differs mainly from that of the other three
species in lacking lateral protuberances and possessing only one pair of
submesial protuberances on the anterior part of sternite XIV, which are
very slender and set relatively far apart. In the other three species the
thelycum bears two pairs of protuberances or often three in S. melantho, and
the submesial ones are usually rounded or oblong, or if slender, roughly fusi-
form and disposed transversely.

Etymology.—This species is named for my brother, Dr. Alfonso Pérez
Farfante, my companion during my first explorations of the seashore. The
specific name is to be treated as a noun in apposition.

VOLUME 94, NUMBER 2 639

Acknowledgments

Thanks are due Horton H. Hobbs, Jr., and Fenner A. Chace, Jr., of the
Smithsonian Institution, and Bruce B. Collette, of the National Marine Fish-
eries Service, Systematics Laboratory, for their criticisms of the manu-
script. Maria M. Diéguez prepared all of the illustrations except those of the
antennular flagella which were drawn by Keiko Hiratsuka Moore; to both
I am most grateful.

Literature Cited

Man, J. G., de. 1907. Diagnoses of new species of macrurous decapod Crustacea from the
‘‘Siboga-Expedition.’’ II.—Not. Leyden Mus. 29:127—147.

Pérez Farfante, I. 1969. Western Atlantic shrimps of the genus Penaeus.—U.S. Fish Wildl.

Serv., Fish. Bull. 67:461-591.

, and H. R. Bullis, Jr. 1973. Western Atlantic shrimps of the genus Solenocera with

description of a new species (Crustacea: Decapoda: Penaeidae).—Smithson. Contrib.

Zool. 153, 33 pp.

, and D. L. Grey. 1980. A new species of Solenocera (Crustacea: Decapoda: Soleno-

ceridae) from northern Australia.—Proc. Biol. Soc. Wash. 93(2):421-434.

Starobogatov, Y. I. 1972. Penaeidae (Crustacea Decapoda) of Tonkin Gulf.—/n Fauna Ton-
kinskogo zaliva i usloviya ee sushchestvovaniya. Issled. Fauny Morey 19(18), Acad.
Sci., USSR, Zool. Inst., Nauka, Lenningrad: 359-415. [Translated by the Office of In-
ternational Fisheries, National Marine Fisheries Service. |

National Marine Fisheries Service, Systematics Laboratory, National
Museum of Natural History, Washington, D.C. 20560.

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(Contents continued from back cover)

A new Goniasterid seastar, Evoplosoma scorpio (Echinodermata: Asteroidea), from the
northeastern Atlantic Maureen E. Downey 561

A redescription of Tubificoides heterochaetus (Michaelsen) (Oligochaeta: Tubificidae)
H.R. Baker 564

The generic status and distribution of Monodella texana Maguire, the only known

North American Thermosbaenacean Jan H. Stock and Glenn Longley 569
Caecidotea salemensis and C. fustis, new subterranean Asellids from the Salem Plateau

(Crustacea: Isopoda: Asellidae) Julian J. Lewis 579
A review of the Platysquilla complex (Crustacea, Stomatopoda, Nannosquillidae),

with the designation of two new genera Raymond B. Manning and David K. Camp _ 591

Anatomy of Diastoma melanioides (Reeve, 1849) with remarks on the systematic
position of the family Diastomatidae (Prosobranchia: Gastropoda)
Richard S. Houbrick 598
Pseudoplumarella echidna, a new species of Primnoid octocoral frem Queensland

(Coelenterata: Octocorallia) Frederick M. Bayer 622
Solenocera alfonso, a new species of shrimp (Penaeoidea: Solenoceridae) from the
Philippines Isabel Pérez Farfante 631

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CONTENTS

The Tanaidae (Crustacea; Tanaidacea) of California, with a key to the world genera

Jurgen Sieg and Richard N. Winn

A redescription of Echinaster modestus Perrier, 1881 (Asteroidea: Spinulosida) from

the eastern Gulf of Mexico and the West Indies

Richard L. Turner and David B. Campbell

The morphology of Tripolydora spinosa Woodwick (Polychaeta: Spionidae): an appli-
cation of the scanning electron microscope to Polychaete systematics

James A. Blake and Keith H. Woodwick

Systematic notes on certain oscines from Panama and adjacent areas (Aves: Passeri-

formes) Storrs L. Olson
Saccogaster melanomycter (Ophidiiformes: Bythitidae), a new fish species from the
Caribbean Daniel M. Cohen
Falsipatellina, a new name for Pseudopatellina Kenawy and Nyir6, 1967 non Haque,
1960 (Foraminiferida) Drew Haman and Richard W. Huddleston
The nature of the variability in the variable seedeater in Panama (Sporophila americana:
Emberizinae) _ Storrs L. Olson
Western Atlantic sea cucumbers of the genus Thyone, with description of two new
species (Echinodermata: Holothuroidea) David L. Pawson and John E. Miller
Two little-known species of Eleutherodactylus (Amphibia: Leptodactylidae) from the
Sierra de la Macarena, Colombia William F. Pyburn and John D. Lynch
The systematic status of Central American frogs confused with Eleutherodactylus
cruentus Jay M. Savage
Pflugella, new name for Tricellaria Pflug, 1965 (Microproblematica), non Fleming, 1828
(Bryozoa) Richard W. Huddleston and Drew Haman

Heteroptera recently collected in the Ray Mountains in Alaska
Richard C. Froeschner and Libby Halpin
Chriolepis vespa, a new species of Gobiid fish from the northeastern Gulf of Mexico
Philip A. Hastings and Stephen A. Bortone
A new genus and a new species of Helminthoglyptid land snails from the Mojave
Desert of California Walter B. Miller
A new species of Aspidosiphon (Sipuncula) from the western Atlantic Ocean
Edward B. Cutler
Psolidium prostratum, new species, from off the east coast of the U.S.A. (Echino-
dermata: Holothuroidea) David L. Pawson and John F. Valentine
A new species of Homochaeta (Oligochaeta: Naididae) from the west coast of Canada
Peter M. Chapman
First description of the male opposum shrimp, Heteromysis bermudensis bermudensis
(Crustacea: Mysidacea) Thomas E. Bowman
Comments on the Taxonomic Status Cr Tommotia Missarzhevsky 1970 (Tommotiidae:
Microproblematica) Richard W. Huddleston
Notes on west African pipefishes (Syngnathidae), with description of Enneacampus,
n. gen. C. E. Dawson
Gonodactylus siamensis, a new Stomatopod Crustacean from Thailand |
Raymond B. Manning and Marjorie L. Reaka
‘A new deep-sea leech, Bathybdella sawyeri, n. gen., n. sp., from thermal vent areas
on the Galapagos Rift Eugene M. Burreson
A new dwarf crayfish from the Pacific versant of Mexico (Decapoda: Cambaridae)
Alejandro Villalobos-Figueroa and Horton H. Hobbs, Jr.
Coralanthura and Sauranthura, two new genera of Anthurideans from northeastern
Australia (Crustacea: Isopoda: Anthuridae) Gary C. B. Poore and Brian Kensley
Ceratocolax mykternastes, new species (Copepoda, Bomolochidae) parasitic in the nasal
sinus of Haemulon sciurus (Pisces, Pomadasyidae) from Belize
Hillary Boyle Cressey
A redescription of Octopus ornatus Gould, 1852 (Octopoda: Cephalopoda) and the status
of Callistoctopus Taki, 1964 Gilbert L. Voss
A revision of the Nettastomatid eel genera Nettastoma and Nettenchelys (Pisces:
Anguilliformes), with descriptions of six new species
David G. Smith, James E. Bohlke, and P. H. J. Castle

(Contents continued inside back cover)

315

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380

391

404

413

421

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445

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